t\vs
HARVARD UNIVERSITY
Library of the
Museum of
Comparative Zoology
us ISSN 0027.4100
bulletin OF THE
Museum of
Comparative
Zoology
A Review of the North American
Fossil Amiid Fishes
JOHN R. BORESKE, JR.
HARVARD UNIVERSITY
CAMBRIDGE, AAASSACHUSEHS, U.S.A.
VOLUME 146, NUMBER 1
18 JANUARY 1974
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(g) The President and Fellow* of Harvard Colleee 1974
A REVIEW OF THE NORTH AMERICAN
FOSSIL AMIID FISHES
JOHN R. BORESKE, JR.i
CONTENTS
Abstract 1
Introduction 2
Acknowledgments 2
Abbreviations 3
A7?Jia calva 3
Nomenclature 3
Ecology 4
Geographic Distribution 4
Pleistocene Occurrences 4
Diagnosis 5
Morphometries 5
Methods 6
General Proportions and Growth 8
Comparisons with Fossil Forms 10
Discussion 17
Meristics 18
Supravertebral Scale Rows 18
Branchiostegal Rays 20
Fin Rays 20
Vertebral Elements 25
Vertebral Column of Amia calva 28
Vertebral Features 28
Vertebral Dimensions 33
Valid North American Fossil Genera and
Species 37
Amia fragosa 37
Amia uintaensis 47
Amia cf. uintaensis 64
Amia scutata 66
Amia cf. scutata 70
Amia cf. calva 72
Amiidae incertae sedis 72
Specimens Removed from the Amiidae 74
Summary and Conclusions 75
Literature Cited 81
Plates 84
Abstract. North American amiid fishes range
from Cretaceous ( Albian ) to Recent. Amiids are
1 Museum of Comparative Zoology, Harvard
University, Cambridge, Massachusetts 02138
Bull. Mus. Comp. Zool, 146(1): 1-87, January, 1974
common fossils in Late Cretaceous and Tertiary
freshwater deposits and apparently occupied a
habitat much like that of the Recent species Amia
calva. Morphometric, meristic, and cranial char-
acters of articulated specimens from the Fort
Union Fonnation (Paleocene), Green River For-
mation (Eocene), Florissant Fonnation (Oligo-
cene). Pawnee Creek Formation (Miocene), and
a Recent A. calva sample from Wisconsin have
been used here in an attempt to revise the taxon-
omy and evolutionary history of the group.
Whereas seven genera and twenty-three species
of fossil amiids have been described on the basis
of disarticulated, often isolated elements, only
three taxa have heen described from complete or
partially complete material. Amia fragosa (Late
Cretaceous to Middle Eocene), A. uintaensis (Pal-
eocene to Early Oligocene), A. scutata (Early to
Middle Oligocene), and A. calva (Middle Plio-
cene to Recent) are here considered the only
valid taxa. Amiid remains are first known in the
North American fossil record from the Early Cre-
taceous (Albian) Paluxy Formation of Texas.
This disarticulated material shows resemblances
both to A»ii« and to the Late Mesozoic European
genera Uroclcs and Arniopsis. Paramiatus gurleyi
(Romer and Fryxell, 1928) from the Green River
Formation of Wyoming is a synonym of A. frag-
osa. Tlie differences between Amia and the large
Early Cenozoic form Protamia are insufficient for
recognition of Protamia as a genus distinct from
Amia. The Eocene and Oligocene forms Protamia
media, Pappichthys medius, P. plicatus, P. scler-
ops, P. laevis, P. symphysis, P. corsonii, Amia
whiteavesiana, and A. macrospondyla are s\monyins
of A. uintaensis; they were based on undiagnostic
cranial and vertebral characters. Morphometric
and meristic similarities indicate that little evi-
dence exists for maintaining separate Oligocene
species A7nia scutata and A. dictyocephala. Amia
exilis is a synonym of Amia scutata; it was based
on undiagnostic vertebral characters. A. scutata
is morphometrically distinguishable from A. calva
only on the basis of a slightly larger head/stan-
1
Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
dard-length ratio. The Eocene ta.xa Amia de-
pressus, A. newherrianus, A. gracilis, and Htjpamia
elegans are nomina diibia.
Comparison of the fossil forms with the Recent
Amia calva suggests the following ta.xonomic and
possible phylogenetic relationships: ( 1 ) Amia frag-
osa survived until the Middle or Late Eocene,
with no phylogenetic affinities with the modern
form; (2) Amia iiintaensis appears to be closer
than Amia fragosa to the ancestry of Amia calva,
which evolved through an intermediate fonn such
as Amia scutata; (3) establishment of the Recent
species Amia calva had begun at least by the be-
ginning of the Pliocene; and (4) diere are simi-
larities in the Paleocene and Eocene amiid fossil
record of North America and Europe.
INTRODUCTION
Aviia is a genus of freshwater fishes that
includes one of two extant representatives
of the holostean level of organization. It
includes a number of species of which only
Amia calva exists today; other forms of
Amia are found in the fossil record, and
extend from the Late Cretaceous to ap-
proximately the Middle Pliocene. This
study is an attempt to determine the taxon-
omic and phylogenetic relationships among
the various species of Amia. It is established
on osteology as well as on morphometric
and meristic data from both Recent and
fossil forms. This data is used to compare
the available features of the fossil forms
with Recent Amia calva and to detennine
the validity of previous descriptions based
on various osteological, morphometric, or
meristic character-states.
Until recently, a major difficulty in inter-
preting the taxonomy of fossil amiids has
been the paucity of articulated specimens.
Five genera and twenty-one species of fossil
forms have been described from disarticu-
lated, often isolated, elements (Table 19);
only two taxa have been described from
articulated specimens: Paramiatus ii^iirleyi
(Romer and Fryxell, 1928) and Amia
scutata (Osborn et al., 1875). Recent works
by Estes (1964) and Estes and Berberian
( 1969 ) , based on disarticulated elements
from the Late Cretaceous Lance and Hell
Creek formations, are the only published
studies of Amia fragosa, although O'Brien
(1969) completed an M.A. thesis on the
osteology of A. frap.osa, describing articu-
lated specimens from the Late Cretaceous
Edmonton Formation of Alberta.
Much more articulated material is now
available and provides more detailed in-
formation on the cranial and postcranial
anatomy of amiids. These specimens have
been useful in this revision of the taxonomy
as well as in the determination of possible
relationships to European and Asian forms.
In an attempt to understand the evolution
and interrelationships of the fossil and Re-
cent amiids, a growth-series study has been
made on a Recent A. calva sample from
Wisconsin, and is compared moi"phometri-
cally and meristically with the fossil forms.
A great number of fossil specimens, includ-
ing the holotypes and paratypes of all North
American amiid species, have been exam-
ined. Several European taxa have been
studied at the British Museum ( Natural
History), London; Museum National d'His-
toire Naturelle, Paris; and the Institut Royal
des Sciences de Belgique, Brussels.
ACKNOWLEDGMENTS
I am especially grateful to Professor
Richard Estes (University of California at
San Diego) for his advice and criticism in
the preparation of this manuscript. Cecile
Janot- Poplin and Sylvie Wenz ( Museum
National d'Histoire Naturelle, Paris), and
Karel Liem ( Museum of Comparative Zo-
ology) read the manuscript and offered
criticisms that substantially improved the
text.
Additional thanks are due to Donald
Baird ( Princeton University ) , Henry Booke
and Bany Cameron (Boston University),
William J. Hlavin (Cleveland Museum of
Natural History), Farish A. Jenkins, Jr.
(Museum of Comparative Zoology),
Charles Meehan ( Chamberlayne College),
Robert R. Miller (University of Michigan),
David Pariis ( New Jersey State Muse-
um), Colin Patterson (British Museum of
Natural History), Clayton Ray (National
Museum of Natural History), Bobb Schaef-
fer (American Musevun of Natural History),
Hans-Peter Schultze ( Geologisch-Paleon-
I
Fossil Amiids • Borcske
tologisches Institiit der Gcorg-Aiigust-Uni-
vcrsitiit, Gottingen), Keith Thomson (Yale
University), and Hainer Zangerl (Field
Museum of Natural History) for their help-
ful suggestions. I am also grateful to Leslie
Whone for preparation of tables, and to Siri
Falck-Pedersen Boreske, Laszlo Meszoly,
and Charles Chamberlain for illustrations.
This study was supported by grants from
Sigma Xi, Marsh Fund, and the Albion
Foundation.
ABBREVIATIONS
AMNH — American Museum of Natural
History, New York, New York.
ANSl^ — Academy of Natural Sciences of
Philadelphia, Philadelphia, Pennsylvania.
BMNH — British Museum (Natural
History), London, England.
CM — Carnegie Museum, Pittsburgh,
Pennsylvania.
F: AM — Frick- American Museum
Collection, New York, New York.
FHKSM— Fort Hays Kansas State Museum,
Hays, Kansas.
FMNH — Field Museum of Natural History,
Chicago, Illinois.
FSM — Florida State Museum, Gainesville,
Florida.
MCZ — Museum of Comparative Zoology,
Harvard University, Cambridge,
Massachusetts.
MNHN — Museum National d'Histoire
Naturelle, Paris, France.
NMC — National Museum of Canada,
Ottawa, Canada.
PU — Museum of Natural History,
Princeton University, Princeton, New
Jersey.
ROM — Royal Ontario Museum, Toronto,
Canada.
SMUSMP— Shuler Museum of Paleontol-
ogy, Southern Methodist University, Dallas,
Texas.
UA — University of Alberta Museum,
Edmonton, Canada.
UCMP — Museum of Paleontology,
University of California, Berkeley,
California.
UMM — West Texas Museum, University of
Texas, El Paso, Texas.
UMMP — l^ni\ersity of Michigan Museum
ot Paleontology, Ann Arbor. Michigan.
UMMZ — University of Michigan Museum
of Zoology, Ann Arbor, Michigan.
USNM— National Museum of Natmal
History, Wa.shington, D.C.
YPM — I'eabody Museum of Natural
History. Yale University, New Haven,
Connecticut.
AMI A CALVA LINNAEUS, 1766
Amid calvii is the only extant species of
the family Amiidae. It is a predaceous fish
that exclusively inhabits fresh waters of
the eastern LTnited States. Except for the
gar, Lepisosteus, Amia calva is the onl\'
other living representative of the holo.stean
fishes. Its common name, "bowfin," refers
to the long dorsal fin that arches in a bow
over most of the length of the fish's back.
Amia calva has previously been known as
the dogfish, marshfish, mudfish, grindle, or
lawyer.
The osteology of Amia calva has been
extensively described and discussed by
Schufeldt (18S5), Bridge (1S77), Allis
(1889, 1897), and Goodrich (1930). The
following discussion is limited only to the
nomenclatural problems, ecology, geo-
graphic distribution, and character-states
of Amia calva that are relevant to study of
the fossil forms.
Nomenclature
Jordan and Evermann (1896) noted that
although Linnaeus (1766) had applied the
binomial name Amia calva to the genus,
Gronow (1763) had earlier used Amia as
a nonbinomial name for fishes presently
classified as Apoiion Lacepede. They fur-
ther suggested that should Gronow's earlier
ipplication of the name be given prece-
dence and transferred to Apof^on, then
Ainiatus Rafinesque (1815) should replace
Amia Linnaeus. Jordon (1906) stated that
this transfer of names was a necessary com-
pliance with the rules of nomenclature, but
later (1919), although citing Opinion 20
Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
( 1910 ) of the International Commission on
Zoological Nomenclature which favored
Gronovv's priority, Jordan found tlie trans-
fer of names inconvenient, for most autliors
had rejected Gronow's names as pre-
Linnaean. In 1925, Jordan recommended to
the Commission that certain names of
Gronow supported by Opinion 20 be re-
jected in favor of the more accepted
Linnaean terminology. The Commission's
Opinion 89 ( 1925 ) resolved ( among others )
the nomenclatural problem of Amia, by con-
curring with Jordan's recommendation that
". . . Amia Gronow be set aside in favor of
Amia Linnaeus, even if other names of
Gronow are allowed." Rafinesque's name
Amiatus is then a junior synonym of Aynia
Linnaeus.
Some later workers seem to have been
unaware of Opinion 89. Thus Hussakof
( 1932 ) accepted the validity of the transfer
of the name A7nia Gronow to the percoid
teleost Apo^on. Romer and Fryxell (1928)
named their fossil amiid from the Eocene
Green River Formation Paramiatus instead
of Paramia, and Whitley ( 1954 ) changed
the name of Lehman's (1951) fossil amiid
from the Eocene of Spitzbergen from Pseu-
damia to Pseudamiatus. The latter is invalid
as Pseudamia was a valid name in itself and
Pseudamiatus is its junior synonym regard-
less of the Amia- Amiatus controversy.
Ecology
Aside from notes regarding breeding, diet,
and zoogeographical occurrences, little has
been written in the past 50 years about the
ecology of Amia calva. Dean (1898) and
Reighard ( 1903 ) have made the only ex-
tensive published investigations of the
habits and habitat of the fish. A thorough
study of the biology of A. calva throughout
its range is long overdue.
Geographic Distribution
The distributional map of Amia calva
( Fig. 1 ) is based on information drawn
from Hubbs and Lagler (1967), and Blair
et al. ( 1968 ) , and from examinations of
unpublished records at the Ohio State Uni-
versity Museum of Zoology, Museum of
Comparative Zoology, and the University
of Michigan Museum of Zoology. The dis-
tribution limit is a flexible boundary allow-
ing for seasonal occurrences and other
natural variations. The known northern
limit of A. calva extends from the Missis-
sippi drainage system in Minnesota south of
Duluth, eastward through Lake Nipissing
and the Ottawa River to the St. Lawrence-
Champlain basin ( encompassing Quebec as
far north as Quebec City, and Vermont).
A. calva is distributed throughout the Great
Lakes region, but is not found in the Lake
Superior drainage basin, except in its outlet,
the St. Mary's River. Southward, it has
been recorded from the Hudson River to
western Connecticut ( recorded as the result
of introduction; Hubbs and Lagler, 1967);
Harrisburg, Pennsylvania, to the Susque-
hanna River; and along the Atlantic slope
to the Carolinas and Florida. Westward,
A. calva occurs along the Gulf Coast to
southern Texas as far south as Brownsville,
and northward, through eastern Texas,
southeastern Oklahoma, northwestern Ar-
kansas, eastern Missouri, and approximately
50 miles west of the Mississippi River to
Brainard, Minnesota.
Pleistocene Occurrences
Amia calva has been reported from only
three Pleistocene localities: (1) Chicago,
Illinois, (2) Vero Beach, Florida, and (3)
Itchtucknee River deposits, Columbia
County, Florida (MCZ 9524, 9529, 9542).
Hay (1911: 552) reported "Professor Frank
Baker (Chicago Academy of Science) has
shown me a considerable part of the skele-
ton and scales of a bowfin which he found
in the Pleistocene clay near Chicago." A
thorough search of the Chicago Academy
of Science collecrions failed to produce this
specimen. Hay (1917, 1923) listed Amia
calva among the fossil vertebrate remains
found in the Pleistocene sands at Vero
Beach, Florida. Swift ( 1968 ) , in his review
of fossil fishes of Florida, figured Hay's
Aiiiia specimens (left dentary and a gular
plate; FSM collections) and concluded that
Fossil Amiids • Borcskc
/ /«°«'m;;; L.._ ; ,
/■ \ r — ■'• ..—■•J^
! \ : NORTH DAKOTA ; ''w
, L . I \ I \minnesota
'""^GO- ^^ > • to ' ■»
^^^^^^■>.-,. / /o* ^1 !
I liJUH—^ m o fe'" oo ^— i
/ / 'l-^-.- i '^
V / ° /'^OlORAdS"— ^ — 1 •(__
\ / /■ j. VMlSbl) I
\ /' ; ; KANSAS ' ' "^
\ / J ! \
\ L .' ^ I o ,
\ .j^^'^oZ. f. ! I
\ I ; "^w MEx-6 T-«- -I
V; • Q *• .OKLAHOMA L
T / -TEXAS ^ 7;
> /• I ^
.. ^ / i '.
Fossil forms of "••v^ ' ! .
© /i3/77/(7 Sp. 'N
• A.fragosa \ s'"~-.
° A.uintaensis ''■^ '\
a A.c\.uintaensis \
' A. scuta to *•
» /J.cf. scuta fa \
■ Amiidae incertae sedis
X Pleistocene location of A.calva
Fig. 1. Distribution of km\a calva. Fossil occurrences of Amia spp. ore represented by symbols explained in the
legend.
A. calva was probably very common in the
Pleistocene fresh waters of the United
States. The pancity of Pleistocene material
does not necessarily mean the fish was not
common in the Pleistocene, but does indi-
cate that A77ua remains have not been
extensively collected or identified in exist-
iii'j; museum Pleistocene collections.
Diagnosis
Vertebral meristics similar to A. sctitata,
but head/standard-length proportion (0.271
mean) is smaller than in the fossil forms.
Extrascapular strap-shaped and relatively
wide at midlitie, as in A. scutata, but pos-
terior edge is curved so that it is proximally
convex, then concave toward the distal
corner, which results in a posterolateral
projection. Pterotic borders frontal pos-
teriorly rather than laterally; anterior end
is as wide as posterior end. Orbital excava-
tion is shallower than in other species, with
a mean depth-to-length ratio of 0.100. In-
fraorbital 4 is smaller than infraorbital 5,
less robust than in fossil Amia. Preopercu-
lum as wide dorsally as ventrally. Symphy-
seal incurx'ing of dentary relatively less than
ill A. fraii^osa, ])ut greater than in A. scutata
and A. uintacnsis: little or no overlapping
of dorsal coronoid articulation surface on
ventral siuface of ramus; deep Meckelian
groov^e. Ventropostcrior process of cleith-
rum less sculptured than in other species of
Amia. V^omerine teeth shaip, conical, num-
bering between 15-27, more anteriorly
placed than in A. uintacnsis or A. fraii.osa.
Bones less ossified than in fossil Amia.
Greatest known standard-length 650 mm.
MORPHOMETRICS
Comparison of morphometric and meris-
tic data of Recent and fossil Amia has
facilitated an e\'aluation of the taxonomy as
well as clarified anatomical trends. Many
6 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
generic or specific character-states for Amia
''dicfyocepJmla,'^ Amia scutata, "Paramiatus
gurleiji,^' and Amia fragosa have been pre-
viously estabhslied on osteological data
based primarily on gross anatomical propor-
tions (head/standard-lcngth ratio and posi-
tions of insertion of pelvic and anal fins/
standard-length ratios) and skull propor-
tions (parietal/frontal and operculum
width/length ratios). Meristic character-
states have also been used for A. "dictijo-
cephala" and A. scutata.
Altliough an age-growth analysis on Amia
calva was done by Cartier and Magnin
(1967), no moiphometric investigation of a
growth-series of Recent A. colva has yet
been completed or used for comparison
with fossil forms. Estes and Berberian
(1969: 10) suggest that knowledge of the
growth-series of A. calva would be of con-
siderable importance in tracing the ancestry
of the modern species.
Hammett and Hammett (1939) made a
moi-phometric study of the Recent Lepisos-
teus platijrhinciis, taking length dimensions
of a sample of live fish from Florida. Since
Lepisosteus, like Amia, is one of the two
extant holosteans, their analysis is poten-
tially useful in providing information on the
ancient species. However, they did not
actually compare the live material or data
with any fossil material.
According to Imbrie (1956), Simpson
et al. (1960), and Gould (1966), growth
studies offer excellent means with which to
clarify evolutionary and taxonomic prob-
lems in the fossil record. An interesting
model utilizing morphometric data for
synonymy of fossil forms was made by
Thomson and Hahn ( 1968 ) on the growth-
series patterns of Devonian rhipidistian
fishes, in which they showed that Thiirsius
clappi was actually a juvenile form of
Eusthenopteron foordi. In studying fossil
material, as Thomson and Hahn (196S:
201) indicate, there is a problem in deter-
mining the age, sexual matmity, and envi-
ronmental regime of the animal. Also, of
course, it is necessary to have sufficient fos-
sil material with which to erect an adequate
growth-series.
This present analysis is undertaken (1)
to determine w hether skull and axial skele-
tal proportions of amiids are isometric or
allometric with increasing size, (2) to
establish the variation in meristic charac-
ters of Recent A. calva, and ( 3 ) to compare
moiphometrics and meristics of Recent A.
calva with those of the fossil forms. This
study utilizes a small sample of 18 Recent
A. calva specimens from the St. Croix River,
Wisconsin. Measurements were taken from
a growth series that includes the size range
of most of the articulated fossil forms. The
largest A. calva specimen, from St. Joseph
County, Michigan (UMMZ 197683), was
analyzed to see whether the large specimen
would agree with the anatomical propor-
tions and meristic characters of the Wiscon-
sin specimens. Three smaller specimens
from Pewaukee, Wisconsin (MCZ 8970),
were also included. The fossil sample con-
tains six complete and ten partially com-
plete amiid specimens ranging in age from
Late Cretaceous to Late Miocene which,
although moiphometrically similar in vary-
ing degree, are too few to warrant conclu-
sions in themselves.
Methods
Measurements chosen for this study ( Fig.
2) are those of Hubbs and Lagler (1967:
20). In fossil forms, because of the lack of
preservation of internal soft anatomy as well
as the impossibility of determining their
interbreeding potential, these particular
measurements necessarily assume an in-
creased taxonomic significance, since they
often provide the only viable parameters
with which to designate genera and species.
Measurements taken on A. calva are limited
to those also represented in the fossil speci-
mens. Each of the A. calva measured was
X-rayed, except for three small specimens,
which were cleared and stained. The range
of error for aj] measmements taken on Re-
cent and fossil material is ±0.04 mm. The
range of error for the ratios is ±0.08 mm;
Fossil Aaiiids • Boreske
Fig. 2. Index to the measurements used, superimposed upon an outline drawing of Amia.
Key for body measurements:
TL = Total-Length
SL =z Standard-Length
H zi: Head-Length
C ^ Caudal-Length
Pf =3 Insertion of Pelvic Fin
P = Insertion of Anal Fin
HL =: Standard-Length minus Head-Length
ML := Standard-Length minus Mandible-Length
Key for abbreviations of cranial elements used in morphometric study:
M
=
Mandible
G
Gular
|5
Infraorbital
F
=
Frontal
Par
=
Parietal
O
zi:
Operculum
Table 1. Length dimensions of 22 specimens of Amia calva L.: 21 from Wisconsin
(MCZ 8970'), 1 from Michigan ( UMMZ 197683)'*
Measurements in mm
Specimen
Class Range
Code
Total Length
No.
TL
SL
ML
H
HL
Pf
P
c
1*
80.0
1
80.0
70.5
57.0
22.0
48.5
32.5
12.5
10.5
2*
95.0-105.0
2
100.0
85.0
70.0
25.0
60.0
39.5
16.0
15.0
3
207.0-212.0
6
210.0
175.0
145.0
50.5
124.5
80.9
35.0
35.0
4
227.9-232.0
4
230.0
193.0
161.0
54.6
138.4
88.5
35.5
36.0
5
241.0
241.0
199.0
165.9
56.8
142.2
93.5
38.0
42.0
6
291.0
291.0
237.0
197.0
64.0
173.0
115.0
52.5
54.0
7
310.0
310.0
248.0
207.0
68.5
179.5
112.0
46.0
62.0
8
339.0
339.0
274.0
230.0
73.0
202.0
125.0
51.0
64.0
9
385.0
385.0
313.0
259.0
82.0
231.0
142.0
52.0
72.0
10
433.0
4.33.0
349.0
293.0
91.0
258.0
170.0
71.0
84.0
11
475.0
475.0
.399.0
335.0
103.0
296.0
181.0
82.0
76.0
12
507.0
507.0
423.0
359.5
109.0
317.0
192.0
93.0
81.0
13»»
756.0
756.0
648.0
545.3
164.0
480.0
299.0
138.0
102.0
See Figure 2 for abbreviations.
8 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
this margin of error is graphically inconse-
quential in this study. Specimens of A. calva
whose total length was between 207 mm
and 507 mm were selected because this
range of A. calva would provide the best
information for comparison with the fossil
species. Twenty-two specimens of A. calva
were measured (Table 1). Eighteen of
these are from the St. Croix River, Wiscon-
sin. These 18 specimens of A. calva fall
into ten categories arranged here by ap-
proximately 20-30-mm class range incre-
ments in total-length. Although these
categories represent arbitrary rather than
biological growth stages, they provide
suflBcient information on the morphologic
size changes of A. calva. Three smaller
specimens (MCZ 8970, also from Wiscon-
sin) witli a size range of 80-105 mm
total length (TL) were included to de-
termine whether they would follow the
predicted allometric effect on the growth-
series Hne, since, as Thomson and Hahn
(1968: 205) note, it is a common feature
for the early stages of juvenile animals to
have heads proportionately larger than the
bodies. Hay (1895) notes that an 80-mm
A. calva is beyond the embryonic stage and
is an early juvenile with most of its bones
at least partially ossified. The 80-mm speci-
men has a proportionately larger head to
standard-length ratio than the other mem-
bers of the growth-series (Table 3). Al-
though this ratio decreases slightly with
increasing size, the head /standard-length
ratio of 0.312 for the 80-mm specimen does
not deviate far from the growth-series line
(Figs. 3-4).
The largest specimen (UMMZ 197683)
was used as a size limit for the other end
of the growth-series continuum. It may be
assumed that this fish had already reached
the size or point of maturity at which fish
normally begin to decrease their rate of
growth. This specimen still retains the
morphological proportions of the smaller
specimens (Figs. 3-4) and, like them, falls
remarkably close to the constant relative
size-growth lines of the various proportions.
Although from Michigan, this specimen
does not appear to deviate from the growth-
series line established by the Wisconsin
specimens of A. calva. The Michigan speci-
men of A. calva, since it agrees with the
growth-series continuum established by the
Wisconsin specimens, is helpful in extend-
ing comparison to the larger fossil amiids:
"Paratniatus gurleiji" (FMNH 2201), Amia
fra^osa (MCZ 5341), and Amia uintaensis
(PU 13865), which are outside the size
range of the Wisconsin sample.
General Proportions and Growth
Allometric growth, according to Gould
(1966: 595), describes geometrically pro-
gressive change in shape or proportions
with size, and is generally represented by a
curvilinear line or, in certain cases, by a
straight line in which the Y-intercept is
significantly different from 0.
For the Amia calva growth series dis-
cussed here, the ordered pairs correspond-
ing to the proportions in each series have
been plotted on a graph, as well as the
straight line corresponding to the equation
y = a + bx (of the best fit computed ac-
Table 2.
Length
DrMENSIONS
; OF 6 ARTICULATED FOSSIL
AMIIDS
Measurements in
mm
TL
SL
ML
H
HL
Pf
P
C
A. scutata PU 10172
'■«yo4.o
339.0
276.5
106.0
233.0
159.0
73.0
e«t65.0
A. scutata UMMP V-57431
—
388.0
313.8
121.0
267.0
183.0
83.0
A. kehreri BMNH P33480
249.0
191.0
160.8
59.2
131.8
89.0
38.5
58.0
"Paramiatus gurleiji"
FMNH 2201
702.0
510.0
430.0
157.0
353.0
<'«f265.0
78.0
192.0
A. fragosa MCZ 5341
575.0
455.0
383.0
142.0
313.0
210.0
75.0
115.0
A. uintaensis FV 13865
848.0
664.0
—
214.0
450.0
288.0
116.0
160.0
See Figure 2 for abbreviations.
Fossil Amhds • Boreske 9
320-
280-
240-
200-
160-
120-
80-
40-
I
40
—I 1 I I I
120 200 280
STANDARD
I I I
360
LENGTH mm
680
440
520
600
Fig. 3. Relative growth-lines of head-length (H), pelvic fin insertion (Pf), and ana! fin insertion (P) plotted arith-
metically against standard-length, for 18 specimens of Recent Ami'o calva (A = MCZ 8970 and H = UMMZ
197683 are included for comparison).
cording to the method of least squares); whicli nearly passes through the origin of
the results of these calculations appear in the graph. The coefficient of correlation is
Figures 3-4. Practically all the ratios in almost equal to 1.0 in each case, an indica-
Figure 3 fall onto straight lines, each of tion that the computed straight line provides
10 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
Table 3. Comparisox of length proportions in 22 specimens of Amio calva with fossil amiids
Specimen
Code
H/SL
Pf/SL
P/SL
1
0.312
0.461
0.177
2
0.294
0.464
0.188
3
0.289
0.462
0.200
4
0.283
0.459
0.184
5
0.285
0.470
0.191
6
0.270
0.485
0.222
7
0.276
0.452
0.185
8
0.266
0.456
0.186
9
0.262
0.455
0.198
10
0.261
0.487
0.203
11
0.258
0.454
0.206
12
0.258
0.454
0.220
13
0.259
0.461
0.213
0.258-0.289''
0.452-0.487*
0.184-0.222*
mean = (0.271)*
mean = (0.463)'
mean = (0.199)*
Oligocene
A. scutata PU
10172
0.313
0.469
0.215
A. scutata UMMP V-57431
0.312
0.472
0.214
Eocene
A. kehreri BMNH P33480
"Paramiatus gurleyi"
FMNH 2201
A. fragosa MCZ 5341
A. uintaensis PU 13865
0.310
0.466
0.201
0.308
*'sto.520
0.153
0.312
0.462
0.165
0.322
0.434
0.175
Range and mean exclude Specimen Codes 1 & 2 (MCZ 8970) and 13 (UMMZ 197683;
a very good fit for the ratio series, and that
the relative growth of these three dimen-
sions is essentially isometric rather than
allometric. The Wisconsin specimens (in-
cluding the 80-105-mm specimens ) and the
larger Michigan specimen all fall close to
the line calculated for each of the three
ratios (Fig. 3). The proportions of head-
length/standard-length, insertion of pelvic
fins/standard-length, and insertion of anal
fins /standard-length are shown in Table 3.
The head/standard-length ratio shows a
slight decrease with increasing size, but
this ratio series nonetheless has a very high
coefficient of correlation for the strength of
the linear relationship (Fig. 4).
The lengths of the mandible, parietal,
frontal, and operculum in Recent A. calva
appear in Table 4, and the proportional
ratios in Table 6. The relative growth rate
of each of these proportions is constant with
X and Y-intercepts of the straight line close
to the origin. The coefficient of correlation
for the variables in each of the proportions
is 0.997, 0.975, and 0.997, respectively ( Fig.
5). Combined, these two factors indicate
constant and therefore isometric relative
size-growth of the compared skull element.
Comparisons with Fossil Forms
Six articulated fossil specimens were
available for measurement and calculation
of head /standard-length and positions of
insertion of pelvic and anal fins/ standard-
length (Tables 2-3). The measurements
taken from the fossil forms are as exact as
conditions allow, although it must be
stressed that varying degrees of crushing
and distortion have occurred in fossilization,
and evaluation of the morphometries should
be qualified with this in mind.
Head /standard-length ratios (Fig. 4).
The fossil forms all show a slightly greater
head/ standard-length ratio than does the
Recent species (Table 3; Fig. 4). A.
uintaensis (PU 13865) is the largest of
Fossil Amiids • Boreske
11
Table 4. Length dimensions of Mandible (M), Gular (C), Frontal (F), Parietal (Par),
Infraorbital ^(I''), and Operculum (O) in 22 specimens of A. calva
Measurements in mm
Operci
ilum
Specimen
Dors. -Vent.
Ant.-Post.
Code
M
G
F
Par
18
(OL)
(OD)
1*
13.5
8.0
10.0
5.0
5.0
8.1
7.5
2*
14.9
9.4
11.4
6.1
5.8
9.0
8.4
3
30.0
19.0
18.0
9.7
12.0
14.0
12.9
4
32.0
21.0
20.0
9.9
13.5
15.0
14.2
5
33.5
20.5
19.0
10.0
14.2
16.0
14.9
6
39.0
26.0
25.0
11.0
17.0
16.9
16.5
7
42.0
28.0
25.5
11.5
18.0
18.1
17.5
8
45.0
27.0
26.2
13.5
21.0
19.8
18.5
9
53.0
31.0
31.0
14.7
22.5
22.8
21.8
10
56.0
32.5
32.2
17.0
26.5
22.6
22.5
11
63.0
39.0
38.4
18.5
30.5
28.1
27.8
12
66.5
42.5
39.0
19.5
31.5
27.8
28.5
13**
102.7
—
60.7
30.0
—
—
o MCZ 8970.
«<» UMMZ 197683.
all the fossil specimens, but nonetheless
has a greater head/ standard-length ratio
than any of the others. The head of this
form is so much more elongated than the
head in A. fra^oso (MCZ 5341), A. kehreri
(BMNH P33480), and "Paramiatus gurleiji"
(FMNH 2201) that it offsets the fact that
its vertebral column includes approximately
20 more vertebrae than do these three forms
(Table 9). Thus, although A. idntaensis
Table 5. Length dimensions of Head (H), Mandiiu^ (M), Gular (G), Frontal (F),
Parietal (Par), Infraorbital ^'{V'), and Operculum (O) in fossil amiids
Measurements in mm
Operculum
Dors.-Vent.
Ant.-Post.
H
M
G
F
Par
F
(OL)
(OD)
A. cf. scutata UCMP 38222
—
65.2
—
46.0
23.0
35.0
—
—
A. scutata PU 10172
106.0
62.5
31.2
35.0
16.0
—
29.0
28.0
A. scutata UMMP V-57431
121.0
74.2
44.3
20.0
29.1
27.9**
A. "dictt/ocephala"
AMNfl 2802
111.5
68.0
38.0
17.0
29.1
32.0
30.0
A. kchrcri BMNH P33480
59.2
30.2
—
20.0
8.4
15.3
20.5
19.0
"Paramiatus fiurletji"
FMNH 2201
157.0
80.0
—
58.0
23.6
25.0
40.0
37.0
A. fra^osa MCZ 5341
142.0
72.0
68.7
56.0
22.8
39.0
36.2
A. fragosa MCZ 9264
80.0
40.0
—
26.0
10.5
18.5
—
—
A. uintaensis PU 13865
214.0
—
— .
88.0
34.0
—
55.0
51.0
"Protamia" mongoliensis
AMNH 6372
—
—
81.0
—
—
—
54.0
52.0
A. uintaensis PU 16236
315.0**
220.0
158.0
160.0
60.0
—
—
95.0
A. fragosa MCZ 9291
— ■
—
—
—
—
—
27.0
25.0
A. fragosa AMNH 9315
—
—
—
—
29.0**
27.0
A. fragosa UA 5450*
—
—
26.0
10.0
—
—
—
A. fragosa UA 5458*
—
—
30.0
12.0
—
—
A. fragosa UA 5480*
—
—
—
20.0
26.0
24.0
•Data from O'Brien (1969).
•• Est.
12 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
240-
220-
200-
EQUATiONof STRAIGHT LINE- M= 7.268 + (0,234) (SLl
u
y
leo-
COEFFICIENT ol CORRELATION = 0.999
ISO-
MEAN ^ -. 0.271
STANDARD DEVIATION 5^ = 0.0117
COEFFICIENT of VARIATION =4.33%
E
E 140.
X
; 120-
a
S 100-
I
^^
60-
xX'
SO'
/^
40'
/^
20-
^x
0.
320 400 460
STANDARD LENGTH mm
y
EQUATION of STRAIGHT LINE - PF = 1 438 + (0.454)(SL)
■^
288 '
240-
COEFFICIENT of CORRELATION .0.995
MEAN f^ '- 0.463
STANDARD DEVIATION |^ -■ 00131
COEFFICIENT of VARIATION ■ 2 83%
y '"
0 y
y
y
y
y
y
192-
y
144.
96-
y
48.
X
0.
X
240 J20 400
STANDARD LENGTH mm
E
E
■z- 120'
EQUATION of STRAIGHT LINE ■■ P = -5.171 +(O.E27)(SL)
COEFFICIENT of CORRELATION = 0.987
MEAN ^ = 0.199
STANDARD DEVIATION^ .0.0135
COEFFICIENT ot VARIATION '6.78 %
240 320 400
STANDARD LENGTH mm
Fossil Ami ids • Boreske 13
has significantly more vertebrae than these
other forms, this feature is not reflected in
a comparison of head/ standard-length ra-
tios ( Table 3 ) . This is also true, to a lesser
extent, in both A. scutata specimens (PU
10172, UMMP V-57431) from the Ohgocene
Florissant Formation; these specimens fall
into the head/ standard-length range of the
three fore-mentioned forms, but like A.
uintaensis, possess vertebral columns having
nearly the same number of centra as those
in A. calva. Thus in themselves the head/
standard-length ratios are of little help in
comparing the fossil forms, but when
coupled with the corresponding lengths of
the vertebral column (based on number
of centra) they are informative. A. uintaen-
sis (PU 13865) and A. scutata (PU 10172,
UMMP V-57431) have relatively elongated
heads; A. kehreri (BMNH P33480), A.
fragosa (MCZ 5341), and "Paramiatiis g,ur-
leyi" (FMNH 2201) have relatively shorter
heads, since the head/standard-length ratio
is less than might otherwise be expected
considering the smaller total-number of
centra (only two-thirds the number of
centra of A. uintaensis, A. scutata, and A.
calva ) . A tentative growth-line ( also calcu-
lated by the best-fit method ) was included
for A. jragosa on the basis of three speci-
mens (Fig. 4). In comparison with the
growth-line of the Recent species (0.271
mean), it reflects the larger head/ standard-
length ratio of A. fragosa (0.310 mean).
The growth-line computed for A. jragosa
is linear and falls near the origin, indicating
that increase in head size/ standard -length
was isometric, as in A. calva.
Fin relations] lips. In the smaller fossil
forms, the ratio of the point of insertion
of the pelvic fin/ standard-length shows little
deviation from the modern species (Table
3; Fig. 4) except for two Eocene specimens,
"Paramiatus gurleyi" (FMNH 2201) and
A. uintaensis (PU 13865), which fall out-
side of the range on either side of tlie size-
growth line. The greater ratio for "Para-
miatus gurleyi," however, is probably the
result of distortion in its preservation. The
length of the pelvic fin insertion/ standard-
length does not appear to be a satisfactory
taxonomic index, distinguishing neither the
fossil forms from one another nor the fossil
forms from the Recent A. calva.
"Paramiatus gurleyi" (FMNH 2201), A.
uintaensis (PU 13865), and A. jragosa
(MCZ 5341) have a relatively shorter
dimension between the anal fin and the end
of the vertebral column than do A. calva,
A. scutata, and A. kehreri (Fig. 31). Any
attempt to inteipret the fossil data for this
ratio is complicated by the fact that con-
siderable overlap with the Recent species
occurs. Both long-bodied (A. scutata) and
short-bodied (A. kehreri) forms fall within
the range of A. calva, while other long-
bodied (A. uintaensis) and short-bodied
(A. jragosa, including "Paramiatus gur-
leyi") fonns fall below the range of the
Recent species (Table 3). Although the
ratio of anal fin/ standard-length may pos-
sibly be useful in distinguishing A. jragosa
(including "Paramiatus gurleyi") from A.
calva, A. scutata, and A. kehreri, it is
not useful in distinguishing either of the
two fossil fonns from one another or from
A. calva. The smaller dimension indicated
by the low ratios (0.153, 0.165) of A.
jragosa is doubtless a reflection of its
shorter axial column. The relatively small
(0.175) ratio for A. uintaensis is probably
in part the result of its longer head, wliich
increases its standard-length in relation to
the other forms; at any rate, the difference
between the A. uintaensis ratio and the
range for Recent A. calva is not very sig-
nificant.
Mandible I head ratios. A comparison of
Fig. 4. Relative growth-lines (broken-solid lines) of head-length, pelvic fin insertion, and anal fin insertion plotted
arithmetically against standard-length for Recent Amia calva (A = MCZ 8970 and ■ = U/AMZ 197683 are included
for comparison) with compared fossil forms: fl = A. hagosa (A. kehreri) BMNH P33480; f- = A. fragosa (Pararr^iafus
gurleyi) FMNH 2201; f'-^ = A. fragosa MCZ 5341; s^ = A. scufafa PU 10172; s^ = A. scutofo UMMP V-57431;
u = A. uintaensis PU 13865. The broken-dotted line is the "best fit" line for available specimens of A. fragosa.
14 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
no
100
90^
tOuATiON o( STRAIGHT LINE ■ M ■-! 010 + (0-636HM)
COEFFICIENT of CORRELATION ^ 0. 997
MEAN ^ ' 0 609
STANDARD DEVIATION ^ ■ 0 0171
COEFFICIENT of VARIATION -■ 2.81 %
55-
50.
/
45-
EQUATION 0) STRAIGHT LINE ^ PAR = 0 278 + (0 479HF) '^ y
COEFFICIENT ol CORRELATION • 0.975 /^ ^
40-
E
MEAN ^ . 0.495 / /"^
E
±
35-
STANDARD deviation""- 0 0333 ^^ /
COEFFICIENT of VARIATION- 6 73% / / "^ V
O
/ /
X
UJ
30-
f' y
~t
/ /
<
/ /
UJ
25-
/
<
V A
S
/ /•'
20-
1!-
10.
/^'
5.
0.
/
30 40 50 60
SO 90 rOO 110 120
FRONTAL LENGTH mm
o-»r
EQUATION of STRAIGHT LINE : OD ■ - 1.559 + (l.079)(&LI
COEFFICIENT of CORRELATION - 0.997
MEAN g2 -0.964
STANDARD DEVIATION ^ - 0.0326
COEFFICIENT of VARIATION- 3.40%
20 30 40
OPERCULUIM LENGTH mni
Fossil Amiids • Boreske
15
Table 6. Cranial proportions in 22 specimens
OF A. calva
Specimen
Code
M/H
Par/F
OD/OL
1
0.614
0.500
0.926
2
0.596
0.535
0.933
3
0.594
0.539
0.921
4
0.586
0.495
0.947
5
0.590
0.526
0.931
6
0.609
0.440
0.976
7
0.613
0.451
0.967
8
0.616
0.515
0.934
9
0.646
0.474
0.956
10
0.615
0.528
0.996
11
0.611
0.482
0.989
12
0.610
0.500
1.025
13
0.626
0.497
0.586-
0.440-
0.921-
0.646*
0.539"
1.025"
mean
mean
mean
= (0.609)"
= (0.495)"
= (0.964)"
■* Range and mean exclude Sjiecimen Codes 1 & 2
(MCZ 8970) and 13 (UMMZ 197683).
the mandible/head ratios of Recent A. calva
with those of the fossil forms ( Table 7; Fig.
5) indicates that the A. scutata and A.
''dictyocephala" (AMNH 2802) ratios are
very close to those of A. calva. The A.
fra^osa .specimens (including "Taramiatus
^urleijr FMNH 2201 and A. kehreri BMNH
P33480) have a mean mandible/ head ratio
of 0.507, which, when compared to the
A. calva mean ratio of 0.609, indicates a
relatively smaller mandible to head size
(Table 7). Unfortunately, A. uintaensis
(PU 13865) cannot be used in this com-
parison, since the mandibles are buried in
matrix. A reconstruction of a disarticulated
A. uintaensis (PU 16236) specimen from
the Late Pal eocene has been made, and its
ratio is approximately 0.693. Thus man-
dible/head proportions may be valid for
distinguishing specimens of A. fra^usa and
A. uintaensis from one another as well as
from A. calva and A. scutata. This ratio,
however, caimot be used as a valid criterion
for distinguishing A. scutata from A. calva.
The 0.693 mandible/ head ratio of A.
uintaeims indicates that this form has the
largest mouth gape of the four valid species.
A tentative growth-line for the mandible/
head-length proportion of A. fra<i^osa, estab-
lished on four specimens, shows that its jaw
is 16 percent smaller than that of A. calva,
and in this respect confirms Romer and
Fryxell's (1928) reconstruction of "Paramia-
tus ^urleiji."
Parietal / frontal ratios. Only articulated
frontals and parietals were measured for
this study. The frontal-length was taken
from the anteriormost extent of the dermal
sculpture to the median point between the
most anterior and posterior extents of the
frontal-parietal suture; the parietals were
also measured by their midline anteropos-
terior length. The parietal/ frontal ratio of
the fossil forms as compared with that of
the Recent A. calva indicates that, in vary-
ing degree, the fossil species have relatively
shorter parietals and longer frontals ( Table
7; Fig. 5). The largest specimen of A.
uintaensis (PU 16236) is mostly disarticu-
lated, but fortunately the skull table is in-
tact. It has the smallest parietal/ frontal
ratio of all the fossil .species, 0.375. The
Eocene A. uintaensis specimen (PU 13865)
has a slightly larger ratio of 0.386, which
is nearly equal to the Edmonton A. jragosa
(UA 5450). All the other available A.
fragosa specimens, including "Paramiatus
gurleyi' ( FMNH 2201 ), have slightly larger
ratios and are quite consistent, ranging only
Fig. 5. Relative growth-lines (broken-solid lines) of mondible-length plotted arithmetically against head-length,
parietal-length plotted arithmetically against frontal-length, and operculum-depth (anteroposteriorly) plotted arith-
metically against operculum-length (dorsoventrally) for Recent Amia calva (A = MCZ 8970 and ^ ^^ '-"^'^'^^ 197683
are included for comparison when element measurements are available) with compared fossil forms: f =^ Amia
fragosa (A. kehreri) BMNH P33480; f- = A. fragosa (Paramiatus gurleyi) FMNH 2201; f* = A. frogosa MCZ 5341;
i^=A. fragosa MCZ 9264; f"' =: A. frogoso UA 5458; f*' = A. fragosa UA 5450; f = A. fragosa AMNH 9315;
f*^ = A. fragosa MCZ 9291; f'^ = A. fragosa UA 5480; si = A. scufaia PU 10172; s- = A. scufaia UMMP V-57431;
s-' = A. scufafa (A. dictyocephala) AMNH 2802; s'=A. cf. scufafa UC 38222; u^ = A. oin/oensis PU 13865; u- =
A. ointoensis PU 16236; m = A. mongo/zensis AMNH 6372. The broken-dotted line is the "best fit" line for available
specimens of Amia fragosa.
16 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
Table 7. Comparison of cranial proportions of recent and fossil amiids
M/H
Par/F
OD/OL
Recent
A. calva ( 18 ) ( Wise. )
Miocene
A. cf. scutata UCMP 38222
Oligocene
A. scutofa PU 10172
A. scufata UMMP V-57431
A. "dictyocephala"
AMMH 2802
Eocene
A. fce/jreh BMNH P33480
"Paramiatus gurleyi"
FMNH 2201
A. /ragosa MCZ 5341
A. /ragosa MCZ 9264
A. uintaensis PU 13865
"Pappichthys" mongoliensis
AMNH 6372
Paleocene
A. uintaensis PU 16236
Cretaceous
A. /ragosa MCZ 9291
A. fragosa AMNH 9315
A. fragosa UA 5450
A. fragosa UA 5458
A. fragosa UA 5480
0.586-0.646
mean = (0.609)
0.590
0.613
0.610
0.510
0.693*
0.440-0.539
mean = (0.495)
0.500
0.457
0.451
0.447
0.420
0.375
0.385
0.400
0.921-1.025
mean = (0.964)
0.965
0.959
0.937
0.927
0.509
0.407»
0.925
0.507
0.408
0.928
0.500
0.404
—
—
0.386
0.927
_^_
___
0.963
0.926
0.931''
0.923
Est
between 0.400-0.408. One of the specific
character-states that Estes and Berberian
(1969: 6) list for A. fragosa is a frontal-
length of approximately 2.8 times the length
of the parietals, which would give a ratio
of 0.357. This figure is smaller than that of
the known articulated forms, including the
specimens from the Edmonton and Will-
wood formations. Although they may have
placed too much emphasis on this specific
character-state, Estes (1964), Janot (1967),
and Estes and Berberian (1969) are justi-
fied in distinguishing A. fragosa from A.
calva on this basis since the ratio of the
fossil form is smaller than that of the Recent
A. calva, whose parietal /frontal proportions
have a mean ratio of 0.495, the frontals
being approximately twice the length of the
parietals. A tentative growth-line for
parietal/ frontal proportions, established on
six specimens of A. fragosa, including
"Paramiatus gurleyi" (FMNH 2201) and
A. kehreri (BMNH P33480), illustrates this
difference between the fossil form and the
Recent species (Fig. 5). A. kehreri dis-
plays slightly larger parietals, with a ratio
of 0.420, but considering the geographic
and temporal differences from the other
A. fragosa specimens, it is remarkably close
in this feature to its North American rela-
tives. The parietal /frontal proportions of
the specimens of A. "dictyocephala" and
A. scutata fall near the lower end of the
0.440-0.539 range of A. calva, and the
Miocene specimen of Amia ( UCMP 38222),
with its ratio of 0.500, is very near the mean
for A. calva. There is thus a definite trend
from the Cretaceous to the Miocene (and
Recent) toward an increase in parietal/
frontal ratio. The A. uintaensis and A.
fragosa specimens have parietal /frontal
ratios smaller than those of A. calva, while
Fossil Amiids • Boreske 17
the A. ^'dictyucepluila" and A. scutata speci-
mens are close to A. calva in this propor-
tion. There is, however, enough intra-
specific variation of parietal /frontal ratios
in the fossil species to cause an interspecific
overlap of the various forms, so that it is
impossible to detennine any definitive limits
between the consecutive fossil species and
A. calva.
Operculum-depth / operculurn-lengtJi ra-
tios. Although the fossil forms have a
slightly narrower operculum-depth relative
to their operculum-length, they all fall
within the operculum ratio range of 0.921-
1.02.5 for A. calva (Fig. 5). Table 6 indi-
cates that with increasing size in A. calva
there may be a trend from a narrower to a
slightly broader operculum. Romer and
Fryxell (1928: 521) describe the operculum
of "Paramiatus <!,urleiji" as being greater
dorsoventrally than anteroposteriorly. Al-
though Cretaceous and Eocene specimens
of A. fragosa have operculum ratios ( 0.923-
0.963) lower than the mean (0.964)
for A. calva, they still fall within the range
(0.921-1.025) of tlie Recent form. Thus the
variation of operculum shape within A. calva
contradicts Hussakof's ( 1932 ) supposition
that the operculum in "Pappichthys" mon-
goliensis (with a ratio of 0.963) is propor-
tionately narrower than that of A. calva, as
well as Estes and Berberian's ( 1969 ) diag-
nosis that A. fragosa has a relatively shorter
operculum-length as compared with height
than A. calva. Janot (1967) was also cau-
tious in assigning taxonomic importance to
the operculum proportions because of the
great variability within the Recent species.
The operculum width /length ratios in tlie
fossil specimens show little taxonomic sig-
nificance, although, as Estes and Berberian
(1969: 7) note, there does appear to have
been a slight temporal trend toward a
broader operculum.
Discussion
The six relative growth proportions
(Figs. 4-5) that were plotted for the A.
calva growth-series remained constant and
therefore isometric. This may be explained
by the fact that these ratios are derived
from external rather than internal dimen-
sions, and, as Gould (1966) points out,
it is usually the internal elements that must
increase at an allometric rate in order to
maintain the external surface area, whose
dimensions may be increasing at an iso-
metric rate (see meristic study). It may
be assumed that the relative growth for
these six proportions also maintained an
isometric rate in the fossil forms, since their
ratios invariably fall near the growth-lines
for the Wisconsin A. calva sample (Figs.
4-5). However, this assumption would
have to be confirmed with an actual
growth-series of the fossil forms.
The comparison of Recent with fossil
forms has also made it possible to determine
the taxonomic value of the skull/ body and
skull proportions. The moiphometric com-
parison of the fossil forms with Recent A.
calva suggests the following taxonomic and
phylogenetic trends :
1. All the fossil forms have slightly longer
heads relative to their standard-length
than does the Recent species ( Fig. 31 ) .
Unless the differences in vertebral meris-
tics are also considered, however, the
morphometric data for this feature are
not useful in comparing the various
fossil forms. A. uintaensis and, to a
lesser extent, A. scutata have relatively
longer heads with a vertebral column of
approximately 85 centra. A. fragosa, on
the other hand, is a short-bodied form
(approximately 65 centra) with a short
head. A. calva has a relatively long ver-
tebral column (81-90 centra) with a
shorter head than the other long-bodied
forms (A. uintaensis and A. scutata).
2. Pelvic fin insertion has no taxonomic sig-
nificance.
3. Anal fin insertion may have minor
ta.xonomic significance for the North
American specimens of A. fragosa wliich
are relatively shorter in this dimension
than in the other .species. There is too
much morphological overlap between
18 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
the species, however, to make this a
useful criterion.
4. A. uintaensis has a relatively longer
mandible/head ratio (0.693) than any
of the other species of Amia, while A.
frafiosa has a smaller ratio (0.507
mean). The mandible/head ratio of A.
scutata (0.604 mean) is close to that of
A. calva ( 0.609 mean ) .
5. There is a trend from the Late Creta-
ceous to Late Miocene in the lengthen-
ing of the parietals in relation to the
frontals. Although it is possible to dis-
cern groups that fall into categories of
smaller and larger ratios (Table 7), in-
terspecific moi-phological overlap makes
it difficult to separate any one of the
fossil species from the others on this
criterion.
6. All the fossil forms have operculum
depth /width proportions that fall into
the lower limits of the A. calva range
(0.921-1.025). These ratios show a
slight temporal trend towards increasing
width, but, while this is perceptible, it
is insufficient to indicate taxonomic sig-
nificance.
These trends suggest possible phylo-
genetic relationships between the various
amiid species. The moiphometric similari-
ties indicate that little evidence exists for
maintaining A. scutata and A. "dictijoce-
phala" as separate species. The Oligocene
A. scutata is distinguishable quantitatively
from A. calva only on the basis of a larger
head/ standard-length ratio, and in this fea-
ture it is intermediate between A. calva and
A. uintaensis. The moi-phometric evidence
indicates similarities between A. fra^,osa
(Cretaceous-Eocene), "Paramiatus ii^iirleiji"
( Eocene ) , and A. kehreri ( Eocene ) . Head/
standard-length ratio is approximately the
same among these three forms; insertion of
anal and pelvic fins /standard-length ratios
shows only minor differences. Mandible/
head size and parietal /frontal ratios are al-
most identical. Of all the .species, A.
uintaensis is the most morphometrically dis-
tinct. It has a relatively greater mandible/
head ratio and a smaller parietal /frontal
ratio than A. fra^osa (Table 7). Even
though it possesses approximately the same
total number of centia as A. calva and A.
scutata (Table 9), it still has a greater
head/ standard-length ratio than the two
latter species. Temporally, there are minor
trends in Amia towards lengthening of the
parietals in relation to the frontals, and in-
creasing operculum width to depth.
MERISTICS
Meristic elements have been used in
species diagnoses of various fossil amiids
by Cope (1875), Osborn et al. (1878),
Romer and Fryxell (1928), and Estes
( 1964 ) . A meristic study of both Recent
and fossil species of Amia was undertaken
to determine tlie relative value of such
diagnoses in the taxonomy of the amiids.
A comparison of the number of supraverte-
bral scale rows, the number of branchio-
stegal rays, and the number of pectoral,
pelvic, anal, dorsal, and caudal fin rays com-
prises the first part of the study, while com-
parative vertebral meristics comprise the
second part.
Supravertebral Scale Rows
Cope (1875) differentiated A. scutata from
A. calva and A. "dictyocephala" on the basis
of A. scutata s (USNM 5374) having seven
and a half longitudinal rows of large scales
above the vertebral column. Cope (1875)
described A. "dictyocephala" (USNM 3992)
as bearing ten to twelve rows of scales
above the vertebral column. A count of the
scale rows between dorsal fin distal pterygi-
ophores and the vertebral column in 20 Re-
cent A. calva (Table 8) gave a range of
seven to nine supravertebral scale rows.
Although the number of scale rows will
vary with the region of the trunk anatomy
from which the count might be taken, Cope
did not designate the point at which he
made his scale row count. Also, his speci-
men WAS so poorly preserved that his count
may have been affected by distortion of
the scales. The only way that a valid com-
parison of all the fonns could be made was
Fossil Amiius • Boreskc
19
TaULK fS. (loMl'AHlSON OF MKHISTIC: ELEMENTS IN HECENT AND FOSSIL NOHTH AMERICAN AMIIDS
Supra-
Branchi-
Pecf oral-Fin
Pelvic-Fin
Anal-Fin
Dorsal-Fin
C"aiidal-Fin
vertebral
ostegal
Lepido-
Lepido-
I.epido-
Lepido-
Lcpido-
Scale Rows
Rays
trichia
trichia
Irichia
trichia
tridiia
Amia calva (20)
7-9
10-13
16-19
7-8
8-11
45-49
23-27
Recent
7.5 av.
11.4 av.
16.8 av.
7.2 av.
10.5 av.
48.0 av.
25.7 av.
A. sciitata
YPM 6243"
USNM 4087*
PU 10172'
7
11
—
7
9
47""'
A. scutata
USNM 5374
7.5
—
—
9
—
A. scutata
YPM 6241
8
9
—
23
A. scutata
UMMP V-57431
7
11
17
7
9
46**
A. "dictijoccphala"
USNM 3992
7.8
—
—
7
9
48»»
—
A. "dicttjoccphaJa"
AMNH 2802
11
—
—
A. "dictijoccphala"
AMNH 2670
9*'
—
—
470 »
A. iiintacnsis
PU 13865
7
16
9
9"*
23
A. uintacnsis
AMNH 785
9«o
—
7
10
—
24
A. fragosa
MCZ 5341
8
12
18
7
8
45
19-20
"Paramiatus gu dcyi"
FMNH 2201
7-8
12
17
8
8
44-45**
19
A. fragosa
UA 5506
10
—
A. fragosa
UA 5425
—
19
" All one specimen.
"o Est.
to take the supravertebral scale row count
of both the USNM 3992 specimen and the
other fossil and Recent amiid specimens at
the same point. In this case, I took all
connts on a vertical line at the level of the
posterior pterygiophore of tlie anal fin. I
connted the nnmber of scale rows in speci-
mens of A. frcif^osa, A. uintaensis, and "Para-
miatus ^urleyi" in addition to those of
Cope's types of A. scutata (USNM 5374)
and A. "dictyocephala" (USNM 3992), as
well as referred specimens of A. scutata;
I then compared them with the supraverte-
bral scale row range in A. calva. The
supravertebral scale rows of fossil Amia
(Table 8) appear to fall within the supra-
vertebral scale row range of Recent A.
calva. Although Cope had described A.
"dictyocephakr (on the basis of USNM
3992) as having 10-12 scale rows, I believe
his count is too high. The supravertebral
scale rows in this and other fossil forms are
difficult to observe for several reasons.
Amiid scales are aligned diagonally to the
vertebral column rather than in parallel,
making it often difficult, particularly in
fossil material, to determine to which diag-
onal column the overlapping scale rows
belong. Also, the scales on the USNM 3992
specimen are broken into many parts, and
Cope may therefore have been counting
partial scales as whole ones. I believe that
I obtained a more reasonable estimate of
the supravertebral scale row count in this
specimen in the following manner: I
measured the average of the anteroventral
20 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
width of complete scales from the abdomi-
nal region (in which the scales are the
same size as in other places in the mid-body
region) and then divided that amount into
the distance between the midpoint of the
vertebral column and the dorsal fin distal
pterygiophore. In this case, the quotient
was 7.8, which is comparable with the
counts of approximately 7-9 in the other
Oligocene specimens and in A. calva ( Table
8). No taxonomic significance can thus be
applied to the number of scale rows above
the vertebral column since counts in Recent
and fossil Amia fall within a' relatively
narrow range.
Branchiostegal Rays
The number of branchiostegal rays was
included in the species diagnosis for A.
"dictyocephalo" (AMNH 2802), in which
Cope ( 1875) counted 12 rays. Osborn et al.
(1878) observed 13 branchiostegal rays in
A. scutata (PU 10172) and Romer and
Fryxell (1928) figured 12 such rays for
"Paramiatus ^urleyi" (FMNH 2201).
O'Brien (1969) counted 10 rays in A.
fragosa (UA 5506) from the Edmonton
Formation. On the basis of disarticulated
material from the Late Cretaceous Lance
Formation, Estes (1964) estimated that A.
fragosa would bear 14 branchiostegal rays,
like the Late Jurassic Sinamia zdanskyi de-
scribed from China by Stensio (1935; see
Liu et al, 1963 for range and distribution).
In the Recent A. calva sample (Table 8),
the number of branchiostegal rays ranges
from 10 to 13; the range among the few
known examples of fossil forms is from 10
to 13, an indication that the number of
branchiostegal rays has remained constant.
Fin Rays
Because of confusing duplication of ter-
minology used for fin description in the
literature, I will use that of Lagler et al.
( 1962 ) for the appendicular skeleton unless
I indicate otherwise.
All fin ray counts on Recent A. calva
were obtained from X-rays of 20 specimens
from Wisconsin and Michigan. The counts
taken from fossil forms are as accurate as
conditions allow, although a number of the
specimens are incomplete or show only
traces of the actual fin. The results of this
study must therefore be considered with
this in mind. I ol^tained these counts from
as close as possible to the internal fin sup-
ports rather than to the segmented and bi-
furcated distal lepidotrichia. There is a
one-to-one correspondence between the
number of lepidotrichia and the number of
pterygiophores in the anal and dorsal fins;
however, this is not the case in the pectoral,
pelvic, and caudal fins, which have more
lepidotrichia than fin supports (Fig. 31).
Pectoral fin. The number of pectoral fin
lepidotrichia has not been previously noted
in any of the original species descriptions
of fossil Amia. There are four .specimens in
which it is possible to make a pectoral fin
ray count (Table 8). A. scutata (1), A.
uintaensis (1), and A. fragosa (2) speci-
mens bear 16 to 18 pectoral lepidotrichia, a
number which is approximately the average
for 20 specimens of Recent A. calva which
displayed from 16 to 19 pectoral fin rays
(Table 8). O'Brien's (1969) analysis of A.
fragosa (Edmonton Formation) does not
include any quantitative comparison of its
pectoral fins with those of A. calva. He
does, however, observe that the pectoral
fins are qualitatively similar in the two
species. The pectoral fins of A. fragosa, A.
scutata, and A. uintaensis thus do not vary
meristically from those of A. calva. Lehman
(1951: 8), in his description of Pseudamia
heintzi from the Eocene of Spitzbergen,
notes that the pectoral fin has 13 complete
nonbifurcating lepidotrichia in the visible
portion of the fossil. This count is different
from that of both Recent and fossil North
American amiids, but as Lehman's plate 3
indicates, this difference may be caused by
matrix that overlies the ventral portion of
the pectoral fin, possibly covering addi-
tional lepidotrichia.
Pelvic fin. The number of lepidotrichia
of the pelvic fin was part of Cope's ( 1875 )
species diagnosis for A. "dictyocephala"
(USNM 3992) and that of Osborn et al.
Fossil Amiids • Boreske 21
(1878) for A. scutota (PU 10172). I
counted the lepidotrichia of these speci-
mens as well as those of on(> additional
Ohgocene specimen and compared them
with my sample of A. calva, which showed
between seven and eight pehic lepido-
trichia (Table 8). Although Osborn et al.
( 1878) counted ten pelvic lepidotrichia, my
recount of their A. scutata specimen (PU
10172) showed only seven (Plate 4). The
bifurcation of the fin rays might have been
inadvertently included in their original
count. The holotype of A. "dictyocepliala"
(USNM 3992) (Fig. 27) showed seven
rather than the six lepidotrichia that Cope
(1875) had diagnosed. A specimen of A.
scutata (UMMP V-57431) (Fig. 27A) also
has seven lepidotrichia; both of these are
within the range of Recent A. calva. Of the
remaining fossil forms, A. fraii^osa and
"Paramiatus ^urleyi" have eight, and A.
uintaensis nine, A. uintacnsis being the only
fossil form not to fall within the range of
Recent A. calva. This difference is insuf-
ficient to demonstrate any taxonomic value,
however, at least until more A. uintaensis
specimens are known.
Anal fin. Anal fin lepidotrichia have
been included in the diagnoses of A. "dic-
tyocephala" and A. scutata (Cope, 1875),
and also in the description of A. .scutata
(Osborn et al, 1878). Each of the original
counts of nine anal rays for each specimen
concurs with my recount and also falls
within the range of eight to eleven for Re-
cent A. calva (Table 8). A. jra<i,osa, "Para-
miatus pMrleyi" and A. uintaensis also fall
within the range of A. calva.
Dorsal fin. Although the number of
lepidotrichia in the dorsal fin has been
mentioned by several authors in their diag-
noses of fossil amiids, it is one of the more
difficult meristic counts to obtain, since a
complete dorsal region of the fossil is
required. Cope's type of A. "dictyocephala"
(USNM 3992) lacks a complete' dorsal fin,
so he counted only the 32 dorsal lepidotri-
chia between the beginning of the dorsal
fin and the posterior lepidotrichia of the
anal fin (Cope, 1875). Osborn et al.
(1878) reported 53 dorsal lepidotrichia for
A. scutata (PU 10172), but this must have
been an estimate, since the posterior por-
tion ot the dorsal fin as well as the entire
caudal fin is missing (Plate 4C). As the
two A. scutata specimens with complete
dorsal fins (AMNM 2670, UMMP V-
57431) have, respectively, 47 and 46 dorsal
lepidotrichia (Table 8), it seems that the
count of Osborn ct al. ( 1878) was high and
that the PU 10172 specimen would prob-
ably have corresponded with the other
Oligocene specimens.
Romer and Fryxell's ( 1928 ) diagnosis for
"Paramiatus fiurleyi" includes a dorsal fin
ray count of 45, which they note as being
slightly fewer than the count for A. calva.
O'Brien's ( 1969 ) discussion of Edmonton
Formation A. fra^osa does not include any
counts of dorsal lepidotrichia, although he
does note that the relative length of the
entire dorsal fin in A. fraiiosa is similar to
that of A. calva. In the A. calva specimens
I studied, the dorsal fin rays ranged be-
tween 45 and 49, the average approximately
48. Romer and Fryxell's diagnosis of "Para-
miatus fiurleyi" as having slightly fewer
dorsal lepidotrichia than A. calva is correct,
but this and all the related fossil forms fall
within the lower range of A. calva (Table
8). The number of dorsal fin rays appears
to have little taxonomic value.
It is interesting that the complete Amia
frafi:osa (MCZ 5341), "Paramiatus <^urleyi"
(FMNH 2201), and A. kehreri (BMNH
P33480) specimens have dorsal fins of
nearly the same length and contain the
same number of lepidotrichia as A. calva,
despite the fact that, on the basis of the
number of vertebrae, these species have a
much shorter body (Table 9). This con-
tributes to a proportional difference in the
body forms of these species, since the dorsal
fin in A. fra<^o.sa (including "Paramiatus
il,urleyi") terminates much closer to the
caudal fin than in A. calva (Plate 1; Fig.
31). However, as Shufeldt (1885) and Hay
(1895) implied, it is very doubtful that the
dorsal fin was fused into a continuous struc-
ture with the caudal fin in some ancestral
22 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
amiid. The Late Mesozoic European forms
of Urocles and Amiopsis have a much ab-
breviated dorsal fin that terminates more
anteriorly than does that of the species of
Amia. For Amiopsis dolloi, an Early Creta-
ceous (Wealden) amiid from Bernissart,
Belgium, Traquair (1911) figured 17 dorsal
fin supports, while Lange (1968) estab-
lishes a specific range of 17-25 for the Eu-
European Upper Jurassic Urocles. The
basis of Shufeldt's (1885: 8.5-86) model for
a primitive amiid with a continuous dorsal-
caudal fin was the presence in Recent Amia
calva specimens of what Shufeldt called a
"series of delicate little bones that continue
the interspinous bones of the dorsal fin as
far as the caudal fin." These five bones he
considered to be the continuation of the
dorsal interneural spines; Hay (1895), in
his discussion of Amia calva, refers to them
as "epural interspinous bones." These small
bones can also be seen in several of the
fossil amiids I have studied, especially A.
scutata (YPM 6241), A. fragosa (UA 5425),
and A. uintaensis (AMNH 785) (Fig. 8).
From these fossil forms, however, it is dif-
cult to determine whether the origin of
these bones is from the dorsal or caudal
neural spines. An examination of the caudal
fin of Urocles lepidotus ( Fig. 6; also
Nybelin, 1963: 506, fig. 17), which is
known to have an abbreviated, more an-
teriorly located dorsal fin, shows that these
epural bones are associated with the caudal
fin, which supports the upper caudal lepi-
dotrichia in much the same manner as the
hypurals in the ventral tail region. A
further indication that these epural inter-
spinous bones are not vestigial dorsal spine
supports is found in Traquair's ( 1911 ) plate
7 of Amiopsis dolloi and his plate 8 of
Amiopsis lata (both species from the Creta-
ceous [Wealden] of Belgium); these plates
show the bones to be clearly associated
with the caudal lepidotrichia (Fig. 7).
i
Fig. 6. Urocles lepidofus MCZ 8300, caudal fin.
Fig. 7. Restoration of Amiopsis dolloi, scales omitted
Early Cretaceous (Wealden), Bernissart, Belgium (after
Traquair, 1911).
Fossil Amiids • Boreske 23
There are no intermediate interspinous
bones between these bones in the caudal
region and those of the much more anteri-
orly situated dorsal spine. The fin of a 648-
mm SL Recent Amia calva (Fig. 10) does,
however, confirm tliat Shuf eldt ( 1885 ) was
correct in stating that the epurals are con-
tinuations of the interneural spines. Figure
10 shows three free interspinous epurals,
with a fourth that is either being fused onto
a neurd spine or is actually a single greatly
elongated neural spine. As only two of
these epurals are attached to lepidotrichia,
there is not a one-to-one correspondence
between the two elements, as in the hy-
purals in the main caudal region.
Caudal fin. With the exception of
Romer and Fryxell's (1928) diagnosis of
"Paramiatus g,urleyi," none of the original
descriptions of fossil Amia include counts of
the caudal fin rays. Although Romer and
Fryxell observed 20 caudal lepidotrichia, a
recount shows only 19 (Fig. 8E). Other
fossil forms tliat also show 19 caudal fin
rays are A. fragosa (UA 5425) from the
Edmonton Formation, A. fragosa (MCZ
5341 ) from the Green River Formation,
and A. kehreri from Messel (Andreae,
1895, plate 1, fig. 23). Another specimen
of A. kehreri from Messel (BMNH P33480)
has 18 lepidotrichia (Plate 2). Traquair's
( 1911 ) plate 7 of three specimens of Amiop-
sis dolloi shows between 15 and 17 caudal
lepidotrichia, while Urocles spp. have a
range between 12-18 caudal lepidotrichia
(Lange, 1968). The only Oligocene speci-
men with a complete caudal fin (YPM
6241) has 23 caudal lepidotrichia; the
Eocene specimens of Amia uintaensis show
23 to 24. Although my sample of Amia
calva displays caudal fins with a range of
23 to 27 lepidotrichia, the number of these
caudal fin rays is skewed toward the higher
limit of the range (Fig. 9). There is thus
a considerable difference between the num-
ber of lepidotrichia in Amia fragosa and
the majority of the A. calva specimens. A.
scutata is, however, within the range of the
Recent species, but occupies the lower
limits of the range.
Thus, of all the meristic elements so far
considered, it appears that the greatest dis-
parity between the fossil forms and the
Recent A. calva is in the number of caudal
fin rays. The number of caudal fin rays
therefore appears to have taxonomic impor-
tance and may have some functional as well
as morphological correspondence to the two
different amiid body types.
As discussed in the preceding section on
dorsal fins, there are two attachment bases
for the caudal lepidotrichia: epural inter-
spinous bones and the hypurals. The
epurals are usually attached to only two or
three of the caudal fin rays, while the
remainder of the lepidotrichia are sup-
ported by the hypurals. Nybelin {in 1963:
488) defines hypurals as "those haemal ele-
ments located to the rear of the emergence
of the caudal artery from the haemal canal"
(trans. Lund, 1967: 210) (Fig. lOB). Lund
(1967: 210) agrees instead with White-
house (1910: 592), who defines hypurals
as "any hypaxial elements that support
caudal fin rays" (Fig. lOA). Lund states
that the sole function of a hypural is to
support a caudal fin ray and therefore the
first hypural would be "the first haemal
spine in rem'ward progression to support a
caudal fin ray and the first ural centrum
is the centrum supporting the first hypural
element." Lund's definition is more practical
for paleontological use. Since there is an
intennediate joint (Figs. 8, 10). the major-
ity of the hypurals are not attached directly
to the urals. However, as Shufeldt (1885)
and Hay ( 1895 ) observed, the posterior-
most seven to nine hypurals are ankylosed
to the corresponding vertebrae ( Fig. IOC ) .
This same co-ossified condition of the last
hypurals is also evident in the fossil fonns,
so that the number of these fused hypurals
has remained constant throughout the evo-
lutionary history of Amia. Also, as Figure
10 shows, the seven or eight anteriormost
hypurals of Recent A. calva have a one-to-
two correspondence with the ventral lepi-
dotrichia. In most of the available fossil
amiid specimens, the ventral caudal portion
is poorly preserved, so that it is difficult to
24 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
Fig. 8. Caudal regions: A, Amia scufaia YPM 6241; B, A. scufafa AMNH 2671; C, A. uintaensis AMNH 785;
D, A. fragosa UA 5425; and E, A. fragosa FMNH 2201.
Fossil Amiids • Boreske 25
u
a>
11 ^n ^li ^5 2r~
Caudal Fin Lepidotrichia no.
TT
Fig. 9. Number of caudal lepidotrichia in 20 speci-
mens of Recent Ami'a calva.
arrive at an accurate count of the total
number of hypurals or to verify whether
this one-to-tvvo relationship exists in all the
amiid fossil forms. The only available fossil
form in which this one-to-two hypural-
lepidotrichia coiTCspondence in the ventral
caudal region can clearly be seen is in A.
scutata (YPM 6241; Fig. 8).
Vertebral Elements
Two regions of the vertebral column, the
trunk and the caudal regions, are defined by
their relationships to the ribs, neural arches,
and haemal arches. The trunk region con-
sists of monospondylous vertebrae that pos-
sess paired basapophyses having gradually
changing angles, dorsal neural facets, and
ventral aortal facets. The number of tnmk
vertebrae in my sample of Ainia calva varies
from 36 to 38. The caudal region consists
of three types of vertebrae, listed from an-
terior to posterior: regular monospondylous
centra bearing neural and haemal arches,
diplospondylous centra bearing neither
neural nor haemal arches (neural and
haemal facets still present ) , and ural centra.
Since the neural and haemal facets are still
present in the diplospondylous centra, there
is no way to differentiate the latter from
the monospondylous type in a disarticulated
state. In my sample of A. calva, the number
of regular caudal monospondylous centra
(24-26) fluctuates by two centra, that of
the diplospondylous caudal centra ( 14-17 )
by three (Table 9).
The posterior caudal region of A. calva
consists of two types of urals: centra with
hypurals attached by a layer of cartilage
(free urals), and centra that are fused
directly onto the hypurals, often lacking the
neural arches (fused urals). When dis-
articulated, the fused urals can often be
distinguished from the free urals, since
part of the hypural usually remains fused
to the ural, extending the posterior articular
surface downward. The nonfused (free)
urals cannot be distinguished in a disarticu-
lated state from the monospondylous or
diplospondylous caudal centra. The num-
ber of urals with fused hypurals is readily
counted, since they are distinguishable
from the remainder of the vertebrae. In
order to identify a free ural, it is necessary
to observe the relationship between the
ural and its conesponding hypural and
lepidotrichia. It is often difficult to make
this distinction between free and fused
urals, since the caudal region is seldom
complete in articulated fossil forms. In A.
calva the number of urals with ankylosed
hypurals ranges between seven and nine.
There are approximately seven principal
urals fused to hypurals, followed by one or
two small additional urals that do not ar-
ticulate with the preceding vertebrae but
lie dorsal to the upturned portion of the
vertebral column. Because it is difficult to
discern these urals in smaller specimens of
A. calva, the count may be slightly biased,
and a comparison of the fossil forms with
the range established for A. calva must be
made with this consideration in mind.
I counted the number of centra between
the anterior dorsal fin pterygiophore and
the posterior anal fin pterygiophore, since
Cope (1875) used the number of central
elements between these points as a specific
character for A. "dictyocephala" (USNM
26 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
Fig. 10. Am'ia calva (648 mm SL) caudal: A, Whitehouse (1910) and Lund's (1967) definition of first ural; B,
Nybelin's (1963) definition of first ural; C, first fused ural.
3992 ) . The range for the number of centra
in this region of Recent A. calva is 33 to 37.
There is considerable variation in total
number of centi'a {i.e., segments) in Recent
A. calva (81-90), which may pose a prob-
lem in comparing specific vertebrae. Thus
in two A. calva, for example, the eightieth
vertebral segment of one individual might
not correspond to the same position in the
vertebral column or even type of centrum
as the eightieth segment of the second indi-
vidual. This should be considered in any
comparisons of several A. calva individuals,
as well as in comparisons of the fossil forms,
which share this variation in vertebral seg-
ments (Table 9). Also, fusion of vertebral
elements may occur in Recent A. calva. In
some specimens, as many as five centra
were found fused together at points
throughout the vertebral column; this con-
dition was present to a lesser degree or
absent in other specimens (Tables 10-12).
These fused centra also occur in the fossil
forms, as in A. uintaensis (YPM 6244). The
actual number of such fused centra can
often be established only by counting ex-
ternal features such as basapophyses, neural
facets, aortal facets, or haemal facets.
Romer and Fryxell's ( 1928 ) study of
"Paramiatus gurleyi" is the only pubHshed
description of a complete articulated fossil
amiid. They distinguished this form from
the Recent species by the supposed pres-
ence of a deeper body, and also noted that
the number of centra was considerably less
than in A. calva. The vertebral column is
completely preserved, so that it is possible
to obtain an accurate count of the vertebrae
(Plate IB). "Paramiatus p,iirleyi" has 67
vertebral segments in contrast to the mean
of 86 in A. calva (Table 9). Osborn et al.
(1878) described A. sciitata (PU 10172) on
the basis of a specimen lacking a caudal
fin (Plate 4). Since the specimen is other-
wise complete, they were able to estimate
that their specimen had 82 vertebral seg-
ments.
Cope (1875) described A. "clictyoce-
Fossil Amiids • Boreske 27
Table 9.
Comparison of
VERTEBRAL
CHARACTERS IN
RECENT AND
FOSSIL AMUDS
Number of
Centra
between
Anterior
Number
Number
Number
Dorsal-Fin
of Mono-
of Diplo-
of Ural
Pterygiophore
Total
Number of
spondylous
spondylous
Centra
and Posterior
Number of
Trunk
Caudal
Caudal
with Fused
Anal-Fin
Centra""
Centra
Centra
Centra
Hypurals
Pterygiophore"
Recent
Amia calva (20)
Wis. & Mich.
81-90
36-38
24-26
14-17
7-9
33-37
mean
mean
mean
mean
mean
mean
= 85.8
= 37.3
= 25.2
= 16.2
= 8.3
= 35.5
Oligocene
A. scutata
PU 10172
83***
36
25
15
yooo
35
A. scutata
UMMP V-57431
81***
36
24***
15***
Y* 00
37
A. "dicttjoccphala"
USNM 3992*
—
—
35
Eocene
"Paramiatus ^urlcyi
•*
FMNH 2201*
67
26
19
16
6
26
Amia uintaensis
PU 13865
85
31
26
21
7
36
Amia uintaensis
AMNH 785
25
20
7
—
A. fragosa
MCZ 5341
65
25
18
15
7
25
A. kehreri
BMNH P33480
62***
24
16
16
6***
24
types.
o --
"" ^ including diplospondylous units (as one),
"o" Est.
pluild" from a specimen (USNM 3992) in
which only the mid-body region was pre-
served. He felt that the number of ver-
tebrae between the anterior dorsal fin
pterygiophore and the posterior anal fin
pterygiophore had ta.xonomic significance.
A comparison of this specimen with Recent
A. calva showed that the v'ertebral count of
this region is essentially the same in both
species. This character is therefore not
useful in distinguishing this species from
the Recent form or in characterizing it as
a specific taxon. The specimens of A.
scutata are within the range of A. calva in
total number of vertebrae as well as in the
number of vertebrae in the vimous cate-
gories (Table 9) . Based on the similarity of
number of vertebrae in A. scutata to that of
A. calva, it appears that the amiid vertebral
column has not changed meristically from
Oligocene to Recent.
Additional data from five undescribed
fossil amiid specimens with relatively com-
plete axial skeletons has been of consider-
able help in estimating vertebral counts
of the fossil forms. A complete specimen
of A. uintaensis from the Green River For-
mation (PU 13865) has a complete axial
skeleton (Plate 3). Interestingly, the total
number of centra (85) does not differ from
that of A. scutata or A. calva (Table 9).
The only variation is in the number of
trunk centra and the number of diplospon-
dylous caudal centra. There are fewer
trunk centra in this specimen of A. uintaen-
sis (31) than in A. scutata, which has a
mean of 36, or in A. calva, \\'hose trunk
centra are a mean of 37. A partially com-
28
Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
plete A. uintaensis specimen (AMNH 785),
also from the Green River Formation, shows
almost the same number of diplospondylous
caudal centra as PU 13865 (20-21 respec-
tively). The lesser number of trunk centra
in both specimens of A. uintaensis is thus
offset by a greater number of diplospondy-
lous caudal centra. In comparing the verte-
bral column of A. uintaensis with that of
A. calva, A. scutata, and A. fragosa, it ap-
pears that although A. uintaensis shares the
same total number of vertebral segments
with A. scutata and A. calva, it does not
conform to their proportional division of
the column into trunk and caudal regions.
A. uintaensis has a trunk/ total-number ver-
tebral ratio of 0.365, while A. fragosa has a
ratio of 0.300 as compared to the A. calva
ratio of 0.440. Three complete specimens
referred to here as A. fragosa ("Paramiatus
gurleyi" FMNH 2022, A. kehreri BMNH
P33480, and A. fragosa MCZ 5341) have
vertebral columns that differ proportion-
ately and meristically from A. calva, A.
scutata, and A. uintaensis. A. fragosa has
significantly fewer centra than the other
fossil forms, with approximately 12 fewer
trunk vertebrae and 8 fewer monospondy-
lous caudal centra. It has approximately
the same number of diplospondylous caudal
centra as A. calva and A. scutata, with the
number of fused hypurals also essentially
the same (Table 9). Thus A. fragosa and
A. uintaensis are meristically distinct from
one another and also from A. scutata and
A. calva, suggesting that these two earlier
forms can be taxonomically separated on
vertebral meristic characters.
VERTEBRAL COLUMN OF AMIA CALVA
The existing taxonomy of many North
American fossil amiids is based primarily
on vertebral characters. Many of the spe-
cies of "Protamia," and the genus itself as
described by Leidy (1873a) from the
Bridger Formation, have been established
solely on height/ width proportions and
length (thickness), shape of the neural
and aortal facets, and various foramina of
isolated vertebrae. Fossil species of Amia
from the Bridger and Cypress Hills forma-
tions have also been defined on character-
states of isolated vertebrae. In order to
analyze this usage, variation in vertebral
character-states of A. calva has been studied.
The axial skeleton of Recent Amia calva
is relatively well known. It is one of the few
modem forms that have diplospondylous
vertebral centra posteriorly, a condition
that, according to Schaelfer (1967), func-
tionally increases the flexibility of the pos-
terior part of the body. Shufeldt (1885)
was one of the first to describe the verte-
brae of Amia, and Hay's ( 1895 ) well-
known work on the vertebral column of
Amia provides a relatively complete and
informative description of the axial skele-
ton, as well as one of the first discussions
of intracolumnar variation of the centra.
Hay observed some gradual changes in
centrum proportions, and in the position of
the neural and aortal facets.
Vertebral Features
Dorsal and ventral facets, basapophyses,
foramina, and ridges on the centra have
been used as diagnostic characters in the
taxonomy of fossil amiids. There are three
types of paired facets on the vertebrae:
dorsal neural facets for the neural arches,
ventral aortal facets for the aortal supports,
and haemal facets for the haemal arches.
Neural facets. The neural facets are
shallow depressions under the neural arch
bases, which in life are filled with cartilage.
Cartilage is present between the centrum
and its associated neural arch. Some speci-
mens of A. calva have much deeper facets,
with a small ossified ridge built up on the
borders. These neural facets occur in pairs
on the dorsal surface of both trunk and
caudal vertebrae, and between the two
facets lies a groove that partially receives
the spinal cord.
According to Hay (1895: 7-9), there is a
marked anteroposterior change in the posi-
tion of the neural facets. He contended
that at the anteriormost end of the vertebral
column the neural arch bases occur be-
tween two vertebrae and rest equally on
Fossil Amiids • Boraske
29
VENTRAL
dUJZ^
odZlL^
28
35
36
37
38
4II>
DORSAL
Fig. 11. Configuration of aortal facets (as) and neural facets (ns) on trunk and anterior caudal vertebrae of
Amio calva (339 mm SL).
both; going posteriorly the.se bases shift
gradually backward. He also observed that
there is a change in the spacing of the
neural iuches; they are close together in the
anterior trinik region and more widely
spaced posteriorly. Hay is correct in regard
to the change in spacing of the neural
arches, but he is not altogether correct in
his description of the change in position
of these arches in relation to the centra.
An examination of the Wisconsin A. calva
sample showed that, after the first few an-
teriormost centra and corresponding neural
arches, the middle of the neural arches is
situated at the juncture between the centra.
This placement continues along the axial
column until the first diplospondylous ver-
tebra occurs. At this point, the next five
to seven neural arches are found aligned to
the middle of each of the corresponding
centra, after which the arches appear to
move forward slightly and correspond ir-
regularly to the vertebral bodies.
The configuration of the neural facets
themselves varies in the trunk region of the
vertebral column of A. calva. The neural
arches in the anterior trunk region are
thicker and wider than those in the more
posterior trunk region which have become
more flattened and elongated. The shape
of the neural facets reflects this trend ( Fig.
11). After the first two ccMitra, the facets
assume an hourglass shape, being narrower
in the middle and broader at each end.
30 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
This can be related to the fact that the
neural arches are situated at the juncture of
two centra so that each neural facet sup-
ports the anterior and posterior halves of
two different neural arches, whose bases
are narrow at the extremity and thick in
the center. Although the neural facets in
any given specimen of A. calva conform to
this general trend, the individual configura-
tion of the facets varies slightly. Given this
variation in shape of the neural facets, it is
useless to attempt characterization of the
vertebral column of any amiid species based
on configuration of neural facets.
Aortal and haemal facets. On the ven-
tral side of the trunk vertebrae are two thin
cartilaginous projections that are located
on either side of the dorsal aortal supports.
When the skeleton is dried, these projec-
tions leave marked depressions, which, like
the neural facets, vary gradually from the
first anterior vertebra to the last few trunk
vertebrae; at this point the aortal facets
coalesce with the basapophyses (Fig. 11).
The point where these two elements are
completely merged marks the termination
of the trunk centra, and the next centrum
is that of the first caudal vertebra. These
structures, which were derived anteriorly
from the basapophyses and aortal facets,
here become the haemal facets.
The first pair of aortal facets is very small
and ovoid. The next few centra bear aortal
facets that, as Hay (1895) also observed,
are circular. The following aortal facets
become successively elongated, until, with
the tenth or twelfth vertebra, these facets
have evolved into a long pair of slits, usu-
ally narrower at the midpoint. Posteriorly,
these slitlike aortal facets remain basically
the same shape until, at the end of the
dorsal trunk region, they merge with the
basapophyses to form haemal facets. Hay
(1895: 54-57) states that the cartilaginous
aortal supports penetrate deeply into the
centra of younger individuals, while in
older specimens they rest superficially on
the centra. The aortal facets are deeper
and more distinct than the neural facets.
Beginning with approximately the tenth
or twelfth vertebra, the slit-shaped aortal
facets are vertically situated on either side
of an indentation that contains the aorta
(Fig. 11). The first four centra have
thicker and shorter supports with relatively
little or no space between them. The aorta
lies ventrally under the basioccipital,
which bears aortal supports whose facets
are of the same shape as the first four centra
(Estes and Berberian, 1969, fig. 2B for
A. frafi^osa). The aortal facets of the first
eight vertebrae are different from all other
trunk vertebrae, whose shape, as mentioned
above, is basically an elongated slit. These
aortal supports are thus helpful in dis-
tinguishing the first eight or so vertebrae
from the remainder of the trunk centra in
disarticulated specimens (Fig. 11).
Haemal facets. The haemal facets,
which contain a cartilaginous layer be-
tween the centrum and the haemal arches,
are nearly rectangular-shaped pairs that do
not vary along the caudal portion of the
vertebral column until the first fused urals.
The furrow or indentation that lies between
the aortal facets in the trunk centra con-
tinues in the caudal region between the
paired haemal facets, although it gradually
decreases in width and depth. Unlike the
neural facets, the haemal facets are out-
lined by an ossified border, which can be
helpful in distinguishing dorsal from ventrid
surfaces in disarticulated caudal vertebrae.
Since the ossified walls are tilted 20
degrees posteriorly to accommodate the
haemal arches, which articulate with the
cartilaginous layer diagonally rather than
laterally, those borders are also useful in
determining the anteroposterior orientation
of the centrum.
Basapophyses. Amia trunk centra are
distinguished from the caudal vertebrae by
their having prominent paired processes,
which ha\e been called transverse pro-
cesses, parapophyses, or diapophyses. I
follow the terminology of Bolk et al. ( 1936),
wherein they designate these structures,
which are the processes for pleural ribs, as
basapophyses ( "basalstiimpfe" ) . The first
centrum often lacks these basapophyses
Fossil Amiids • Boreske 31
20
44
24
48
28
56
8
32
<k^
64
36
68
40
Fig. 12. Shape of selected trunk and caudal vertebrae in Amia calva (339 mm SL).
78
(Tables 10-12), which are always present
on the succeeding centra and progressively
become longer until appro.ximately the
twelfth (Fig. 12). The basapophyses are
approximately the same length between
the twelfth and tlie thirty-second centra,
from which point they begin to diminish
gradually in length until the last trunk
centrum, where they coalesce with the
aortal facets. The lengths of the basapo-
physes were not individually measured;
this data would be of little practical use in
a comparison of Recent and fossil material
since these relatively fragile structures are
rarely preserved intact in fossils. The distal
end of each basapophysis is attached to a
pleural rib by means of cartilage. The
proximal ends of the basapophyses are
ankylosed to the ventral half of the verte-
bral body. These paired processes are solid
cyHnders (hollow at the tips) that are
slightly Battened dorsoventrally. Each pair
of basapophyses may not always be of equal
length or diameter, but they are extremely
regular in position. They form two con-
tinuous and symmetrical lines that gradu-
ally come closer together until the last
trunk centrum, where they are separated
only by aortal supports.
An important aspect of the basapophyses
in A. calva is the angle between each indi-
vidual pair which gradually decreases pos-
teriorly. Since the angl(> ])etween the
basapophyses is generally still available in
32 Bulletin Museum, of Comparative Zoology, Vol. 146, No. 1
fossil forms, even in those with broken basa-
pophyses, it is used here as a basis of
comparison between the Recent and fossi!
forms. Since the angles steadily decrease
posteriorly along the vertebral column
(Figs. 12, 14), they are also useful in
orienting disarticulated centra to approxi-
mate position along the column. Although
there is individual variation in these angles
(Tables 10-12), they are nevertheless con-
sistent enough to help in determining the
general position in the column of any
single trunk centrum. The range of angles
extends from approximately 180 degrees
anteriorly to 45 degrees posteriorly. Since
the three A. calva specimens studied were
of varying sizes ( 193 mm SL, 382 mm SL,
and 423 mm SL), it would appear that there
is no significant change in the angles with
increasing size or age of the fish (Fig. 14).
Although this transition is not perfectly
linear, the angles are always decreasing
posteriorly, and at least in the specimens I
measured, there was never an instance of
an angle's measurement being greater than
that of the preceding centrum. The angle
decrease occurs at a fairly constant rate
until approximately the thirtieth trunk ver-
tebra, at which point the rate of decrease
of the angles is much accelerated ( Fig. 14 ) .
The angle of the basapophyses is thus a
reliable parameter in identifying the gen-
eral position of isolated trunk centra.
Foramina, bone ridges, and first centrum.
The trunk centra of A. calva have lateral
foramina that, although lacking the uni-
formity of the neural and aortal facets,
occur in irregular, distinct paired linear
patterns. The foramina of the tnmk and
caudal vertebrae transmit numerous small
blood vessels.
On the lateral surfaces perpendicular to
the anterior and posterior articular surfaces
of the individual centra are prominent bone
ridges. These bone ridges add support to
the arch anlagen, and also help unify the
anlagen into a sturdy, functional vertebral
body (Schaeffer, 1967). Externally, these
bone ridges are not as regular as they are
internally, although they still lie antero-
posteriorly in the lateral and ventral regions
and extend vertically along the basapo-
physes. They are also quite prominent in
the notochordal furrow. Such bone ridges
are not a unique feature of A. calva, and are
common in teleosts.
The centra in A. calva are amphicoelous.
The first four to six centra differ from all
corresponding centra by having the anterior
articular surface more convex than concave.
The first centrum in nearly all specimens
observed lacked basapophyses, and should
therefore be considered a minor taxonomic
character since first centra do occasionally
occur with very small basapophyses. The
Angle of
basapophyses
Fig. 13. Index to the measurements used, superimposed
upon an outline drawing of an Amia calva vertebra.
Fossil Amiids • Boreske 33
^. co/yo verfebrol lengths
1( 24 n <0 <l Si (4 72 10 II
Verl. no.
A.calvo bosopophyseol angles
12 It 21
Vert. ■•.
---^
12 41 41
Vtrt. ■•.
Vx
y
v^i
•-v.'
A. uinfaensis
NEIGIT
mom
I i ii H W
—» — jr
Vtrt. M.
Ti n w
Fig. 14. Intracolumnar variation in the angle of basapophyses, length, height, and width of vertebrae in Re-
cent Ami'o co/vo (A = 423 mm SL; B ^ 382 mm SL; C =: 193 mm SL). Intracolumnar variation in height and width
of trunk and caudal vertebrae in A. uin/oensi's. Vertebral column model based on first six centra from PL) 10101
and fifty-nine centra from CM 25362; missing caudal centra have been interpolated and inferred based on PL) 13865.
The first anterior centra (PL) 10101) were larger specimens and thus the anterior region of the trunk vertebral column
model is "out-of-phase." Vertical lines = last trunk centrum.
ovoid shape of the aortal facets is a constant
feature of all first four to six centra ob-
served.
Vertebral Dimensions
A superficial but often-used character for
diagnosing fossil amiid species has been the
shape of the centrum. Descriptions for
Amia whiteavesiana, A. macrospondyla, A.
exilis, A. elegans, A. depressus, A. newher-
rianus, Protamia symphysis, P. media, P.
gracilis, P. uintaensis, P. plicatus, P. cor-
sonii, and P. laevis include centrum mea-
surements for height, width, and length
(thickness), as well as qualitative descrip-
tions of the fonii and proportions of the
centrum. Because isolated amiid vertebrae
have often been the only anatomical mate-
rial found in the fossil record, the original
diagnoses were obviovisly limited in that a
great deal of emphasis was placed on the
vertebral centrum. In considering a single
centrum shape as diagnostic for an amiid
species, early authors implicitly assumed
the vertebral column to be static, with no
physical change or variation among the
centra other than regional. Many new
species were therefore described solely on
variation in shape from other known amiid
types.
34 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
Table 10. iNiTtAcoLUMNAR variation in height, width, length, and angle of basapophyses
OF VERTEBRAE IN RECENT Amiu colva ( 193 mm SL)
Vert.
Cent.
Height
(mm)
Width
(mm )
Length
(mm)
Angle of
Basapophyses
( Degrees )
Vert.
Cent.
Height
(mm)
Width
(mm)
Length
(mm)
1
3.55
4.35
1.40
44
2.75
2.75
1.35
2
3.65
4.20
1.70
176
46
2.60
2.65
1.15
4
3.70
4.25
1.65
159
48
2.65
2.65
.90
6
3.40
4.15
1.70
159
50
2.55
2.60
1.30
8
3.25
3.90
2.05
153
52
2.50
2.45
1.35
10
3.15
3.65
2.00
145
54
2.50
2.50
1.20
12
3.00
3.65
1.90
140
56
2.45
2.45
1.15
14
3.05
3.60
2.00
138
58
2.50
2.45
1.20
16
2.95
3.55
2.20
134
60
2.30
2.40
1.15
18
3.00
3.55
2.30
130
62
2.40
2.30
1.15
20
3.05
3.40
2.10
122
64
2.30
2.20
1.15
22
2.90
3.65
2.25
118
66
2.30
2.05
1.10
24
2.75
3.25
2.10
103
68
2.25
2.15
1.00
26
3.00
3.50
2.15
99
70
2.30
2.15
.80
28
3.05
3.35
2.20
93
72
2.30
2.10
.85
30
3.00
2.90
2.25
91
74
2.25
2.05
1.00
32
2.95
3.05
2.15
90
76
2.05
1.90
.90
34
3.00
3.00
2.05
74
77
1.87
1.75
.90
36
3.10
3.08
2.25
63
80
1.65
.80'
37
3.15
3.00
2.15
44
82
1.50
.68
38
3.25
3.00
2.10
84
1.25
.53
40
2.80
2.90
2.25
86
.80
.50
42
2.75
2.90
1.50
" Fused.
Table 11. Intracolumnar variation in height, vi^idth, length, and angle of basapophyses
OF vertebrae in recent Amia calva (382 mm SL)
Vert.
Cent.
Height
(mm)
Width
(mm)
Length
(mm)
Angle of
Basapophyses
( Degrees )
Vert.
Cent.
Height
(mm)
Width
(mm)
Length
(mm)
1
6.30
7.21
1.90
180
42
5.50
5.05
3.50
2
6.25
7.41
2.45
173
44
5.15
4.65
2.40
4
6.00
7.50
2.50
161
46
5.00
4.80
2.35
6
5.95
7.25
2.85
161
48
4.90
4.60
2.37
8
5.82
7.10
2.95
159
50
4.70
4.45
2.50
10
5.56
6.90
3.05
154
52
4.70
4.45
1.90
12
5.50
6.65
3.45
143
54
4.65
4.55
2.00
14
5.50
6.25
3.45
140
56
4.70
4.45
2.10
16
5.50
6.25
3.45
133
58
4.80
4.10
1.90'
18
5.35
6.30
3.50
125
60
4.40
4.45
2.00
20
5.25
6.25
3.55
120
62
4.80
4.45
2.12'
22
5.30
6.25
3.55
116
64
4.50
4.10
2.12'
24
5.30
6.25
3.65
113
66
4.40
3.85
2.00
26
5.20
6.20
3.85
110
68
4.00
3.75
2.00
28
5.35
6.15
3.70
110
70
4.05
3.70
1.87
30
5.25
6.00
3.60
105
72
3.75
3.50
1.75
32
5.35
5.80
3.85
100
74
3.50
3.35
1.50
34
5.45
5.50
3.90
92
76
3.20
3.15
1.47
36
5.50
5.15
3.50
78
78
2.65
1.65
38
5.75
5.10
3.55
66
80
2.00
1.40
39
5.70
5.07
3.60
46
82
1.95
1.20
40
5.65
5.05
3.65
84
1.80
1.12
• Fused.
Fossil Amiids • Boreske 35
Table 12. Intracolumnar variation in height, width, length, and angle of basapophyses
OF vertebrae in recent Amia calva (423 mm SL)
Vert.
Cent.
Height
(mm)
Width
(mm)
Length
(mm)
Angle of
Basapophyses
( Degrees )
Vert.
Cent.
Height
(mm )
Width
(mm)
Length
(mm)
1
8.25
9.25
2.80
44
6.45
6.40
3.45
2
8.00
9.15
3.35
172
46
6.40
6.21
3.05
4
7.80
9.05
3.40
156
48
6.20
6.00
3.05
6
7.55
9.20
3.45
150
50
6.10
5.85
3.85
8
7.45
8.80
3.45
142
52
6.25
5.85
3.80
10
7.40
8.80
3.75
140
54
6.10
5.85
2.95
12
7.25
8.75
3.85
132
56
5.95
5.90
2.70
14
7.25
8.75
4.00
126
58
5.80
5.85
2.65
16
7.35
8.70
4.05
119
60
5.80
5.65
2.60
18
7.15
8.50
4.15
117
62
5.65
5.70
2.80
20
7.15
8.50
4.10
114
64
5.60
5.30
2.70
22
7.15
8.40
4.00
112
66
5.55
5.20
2.45
24
7.35
8.35
4.37
103
68
5.50
5.05
2.50
26
7.45
8.30
4.75
103
70
5.30
4.95
2.35
28
7.30
8.20
4.75
95
72
5.30
4.90
2.20
30
7.47
8.10
4.70
90
74
5.20
4.80
2.10
32
7.25
7.95
4.95
90
76
5.05
4.65
1.85
34
7.35
7.45
5.00
67
78
5.00
4.50
2.10*
36
7.35
6.90
5.00
52
80
4.75
4.45
2.25
37
7.50
6.70
4.95
46
82
4.15
1.55
38
7.65
6.50
4.90
84
3.25
1.50
40
7.60
6.50
4.50
86
2.50
1.50
42
7.40
6.20
4.35
88
2.00
1.45
<• Fused.
One of the detailed studies on intra-
columnar vertebral variation is Hoffstetter
and Case's ( 1969 ) work on the vertebral
column of snakes. Measuring individual
centra in sequence, they plotted this varia-
tion; similar graphs are used here (Fig.
14 ) . Three specimens of A. calva ( 193
mm SL, 382 mm SL, and 423 mm SL ) were
dissected and the individual vertebral
dimensions measured to determine verte-
bral variation.
Length. Centrum length (thickness)
was measured anteroposteriorly at the mid-
line, above the basapophyses (Fig. 13).
It was necessary to establish such a control
for this measurement because of the varia-
tion in thickness within each centrum. In
the trunk region, the centra are thickest
ventrally at the neural and dorsal facets.
The caudal vertebrae follow a similar pat-
tern, being slightly thicker ventrally and
dorsally, and thinner laterally. Every
second centrum was measured for length
(Tables 10-12). There is a distinct intra-
columnar variation for this measurement,
although the difference in length between
consecutive vertebrae is usually small.
There is also a general, if somewhat ir-
regular, pattern in centrum length in A.
calva (Fig. 14). The first two or three ver-
tebrae of each A. calva specimen are rela-
tively thin. These are followed by centra
that gradually increase in length until ap-
proximately the last trunk centrum at the
midbody. At this point there is a general
trend again towards thinner vertebrae, al-
though this pattern is erratic, particularly
between the fiftieth and the fifty-fourth
centra, where the thickness is suddenly in-
creased and then decreases. This sudden
change here in vertebral thickness occurs
directly above the midline of the anal fin.
The shortest vertebrae are the fused urals.
Height. Centrum height was taken dor-
soventrally at the midline, between the
aortal and neural facets (Fig. 13). Every
second centrum was measured up to the
fused urals, in which an accurate measure-
36
Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
Table 13. Intracolumnar variation in height, width, length, and angle of basapophyses
OF VERTEBRAE IN Atnla uintaensis CM 25362
Vert.
Cent.
Height
(mm)
Width
(mm )
Length
( mm )
Angle of
Basapophyses
( Degrees )
Vert.
Cent.
Height
(mm)
Width
( mm )
Length
( mm )
7
24.5
32.8
9.8
179
37
19.5
19.5
7.5
8
25.0
32.5
9.8
178
38
19.8
18.8
7.0
9
26.8
32.0
9.5
177
39
19.8
18.5
6.9
10
26.5
32.1
9.5
176
40
20.0
17.5
6.5
11
25.5
32.8
9.0
174
41
—
6.5
12
26.0
31.0
9.1
172
42
19.5
18.0
6.5
13
25.4
31.1
9.5
164
43
18.5
17.5
6.0
14
27.5
31.0
9.5
160
44
18.5
17.5
6.2
15
25.5
30.0
10.0
156.5
45
17.5
17.5
6.0
16
26.0
30.5
9.8
154
46
17.2
16.0
6.0
17
26.5
31.5
9.5
153
47
17.0
16.0
6.0
18
27.0
30.5
9.8
149
48
16.5
16.0
5.5
19
25.1
29.5
9.5
143
49
16.5
—
6.0
20
24.5
29.5
9.5
132
50
16.2
15.5
5.5
21
24.0
30.0
9.5
122
51
16.0
15.0
5.8
22
24.5
28.2
9.0
117
52
16.0
15.5
5.0
23
24.0
29.0
9.5
110
53
15.5
14.0
5.5
24
24.0
28.5
9.0
102
54
15.2
14.5
5.2
25
24.0
26.5
8.5
102
55
15.5
13.2
6.0
26
24.5
27.0
10.0
97
56
15.0
12.2
5.5
27
23.5
25.0
11.0
90
57
14.5
12.0
6.5
28
23.5
25.0
11.0
83
58
14.5
12.5
5.5
29
23.2
25.0
12.0
80
59
14.0
11.5
5.2
30
23.0
22.5
11.0
62
60
14.0
5.0
31
25.0
22.5
11.0
46
61
13.5
11.5
5.0
32
—
11.0
62
13.5
10.5
4.5
33
—
10.0
63
4.5
34
23.0
19.5
9.5
64
12.0
9.0
4.5
35
22.5
9.5
65
8.5
7.0
4.2
36
21.0
20.0
7.5
ment would be obscured by the fusion of
the hypurals. This series of measurements
shows a basic pattern that is similar for
each of the individuals studied, although
there is less intracolumnar variation in the
height than in the length measurements
(Tables 10-12). The greatest height gen-
erally occurs at the anteriormost region of
the column, then decreases slightly until
the midtrunk region (Fig. 14). At this
point the height gradually increases until
it peaks at the end of the trunk region and
the beginning of the caudal section, after
which it decreases again toward the caudal
region (Fig. 14).
Width. The width measurements were
taken perpendicular to the height measure-
ments, at the widest section of the centrum
(Fig. 13). This dimension has a greater
linear slope than the length and height
dimensions ( Fig. 14 ) , which follow a more
bell-shaped curve. There is a greater varia-
tion within the vertebral column for width
dimensions (Fig. 14), as comparison of the
height and width slopes reveals. The great-
est intracolumnar width is always at the
anteriormost portion of the trunk region,
after which this dimension gradually de-
creases. There appear to be two areas
where the rate of decrease is greater, these
being at the terminus of the trunk region
and at the first fused ural.
Height/ width ratio. The centrum height/
width ratio has been a commonly used diag-
nostic character in amiid taxonomy. Hay
(1895: 7) correctly noted that the trunk
vertebrae are somewhat broader than high
( Fig. 14 ) , and at the terminus of the trunk
Fossil Amiids • Boreske
37
region the centra are nearly circular. The
proportions tend to be reversed in tlie
eandal region, however, with the height
generally exceeding the width, although to
a lesser degree than the proportional dif-
ference in the trunk region. The basic trend
in shape through the vertebral column is
tluis a marked horizontally elliptical cen-
trum approaching a progressively circular
one, which then again becomes slightly
vertically elliptical. Thus there is quite a
variation in the centrum shape throughout
the axiiil column, so that no one shape or
ratio of dimensions could reasonably be
considered diagnostic for all the centra of
the vertebral column.
VALID NORTH AMERICAN FOSSIL
GENERA AND SPECIES
Amia fragosa (Jordan, 1927)
Kitidleia fragosa Jordan, 1927: 145.
SUjlomylcodon lacus Ru.ssell, 1928a: 103.
Paramiatus giirleyi Ronier and Fryxell, 1928: 519.
Holotype. NMC 8533e, anterior portion
of right dentary.
Paratypes. NMC 8534a-d, f-n. (a), left
operculum fragment; ( b ) , cranial fragment;
(c), anterior portion of left dentary; (d),
anterior portion of right dentary; (f-g),
coronoid with two styliform teeth preserved;
(h-i), vomer without teeth preserved; (j),
styHform tooth fragment; (k-1), posterior
portion of right dentary; ( m ) , left maxilla;
(n), anterior portion of right dentary.
Ttjpe locality and horizon. Rumsey, Al-
berta. East half of section 31, T 34 S, R 21
W, Rumsey Quadrangle, Alberta; Edmon-
ton Formation.
Age range. Campanian (Late Creta-
ceous) to Bridgerian (Middle Eocene).
Hypodiii^m. Cretaceous. Oldman For-
mation, Alberta: AMNH 5934, palatal frag-
ments with styliform teeth; AMNH 5935,
operculum and dentary. "Mesaverde" For-
mation, Wyoming: AMNH 5932, dentary
and numerous coronoid teeth; AMNH 5933,
vertebrae. Judith River Formation, Mon-
tana: AMNH 10109, left vomer bearing
styHform teeth; AMNH 10110, dentary frag-
ments, vertebrae, and skull elements. Ed-
monton Formation, Alberta: ROM .3064,
coronoid teeth; ROM 3065, dentaries, verte-
brae, and cranial fragments; UA 5398-5507,
articulated and disarticulated specimens
(see O'Brien, 1969 for identifications).
Lance Formation, Wyoming: AMNH 9316,
pterotic; AMNH 9315, operculum; CM
25363, dentaries; PU 17013, dentaries;
UCMP 54013-54015, 54017, 54019, 54021-
54030, 54035-54038, 54040-54056, 54059-
54069, 54070-54120, 54141-54167, 54174-
54180, 54188-54198, 54260, 54262, dis-
articulated elements (see Estes, 1964 for
identifications). Hell Creek Formation,
Montana: PU 17016, 17048, dentaries; PU
17014, coronoid teeth; PU 20554, dentary
and vertebrae; MCZ 9286-9293, 9390-9432,
9559, disarticulated elements ( see Estes and
Berberian, 1969 for identifications ) .
Paleocene. Fort Union Formation, Wy-
oming: PU 17115, coronoid teeth; PU 17126,
coronoid teeth and vertebrae; PU 17117,
dentary and maxilla; PU 21525, portion of
cranial roof with associated dentaries; PU
20523, dentary and coronoid teeth; PU
21174, vertebrae. Paskapoo Formation, Al-
berta: UA 131, dentary, numerous tooth
plates, and vertebrae. Tongue River For-
mation, Montana: PU 20577, vertebrae,
premaxillary fragment, and coronoid teeth;
PU 20578, basioccipital and vertebrae; PU
17068, vertebra and dentary fragment.
Melville Formation, Montana: AMNH 2635,
cranial elements and associated dentaries.
Tullock Formation, Montana: PU 17069,
vomers.
Eocene. Will wood Formation, Wyo-
ming: MCZ 9264, nearly complete skull;
PU 18780, tooth plate; PU 21175, dentary
fragment and coronoid teeth; PU 16756,
dentary and cranial fragments; PU 17649,
anterior portion of skull; PU 21173, skull
fragments and vertebrae; PU 13261-13262,
cranial fragments and coronoid tc>eth.
Golden Valley Formation, North Dakota:
PU 18567, coronoid teeth and vertebrae.
Wasatch Formation, Wyoming: PU 13260,
tooth plates; PU 13259, cranial fragments
and dentaries. Bridger Formation Wyo-
B
Fig. 15. A, Amia calva. Recent, Wisconsin; above, lateral, and below, dorsal views of skull. B, Amia scutafa. Early
and Middle Oligocene; above, lateral, and below, dorsal views of skull (sensory canal system and pit-lines are not
known since skull elements are in articulation). C, Amio fragosa, Late Cretaceous to Middle Eocene; above, lateral,
and below, dorsal views of skull (sensory canal system and pit-lines after Estes, 1964). D, Amia uinfaensis, Paleocene
to Early Oligocene; above, lateral, and below, dorsal views of skull (sensory canal system is only known in the
mandible, operculum, nasal, lacrimal, antorbital, extrascapular, and suprascapular, all of which conform with those
of A. calva).
Abbreviations: a, angular; ao, antorbital; br, branchiostegal rays; d, dentary; ds, dermosphenotic; es, extrascapular;
fr, frontal; io, interoperculum; io- io'' io'* io'^', infraorbital series (suborbitals & postorbifals); la, lacrimal; m,
maxilla; n, nasal; op, operculum; p, preoperculum; pa, parietal; pt, pterotic; r, rostral (ethmoid); s, suprascapular; so,
surangular; sm, supramaxilla; so, suboperculum. Dotted lines indicate the sensory canal system; dashed lines indi-
cate pit-lines.
38
Fossil Amiids • Boreske
39
ining: YPM 6245, vomer and cranial frag-
mcMiVs; YPM 6246, vertebrae; YPM 6247,
dentary; YPM 6248, vertebra and cranial
fragments; YPM 6254, verte]:)rae, basioccip-
ital, vomer; YPM 6261, left opercnlnm;
ANSP 5630, vertebra. Green RivcT Forma-
tion, Wyoming: MCZ 5341, FMNH 2201,
complete specimens.
Known distribution. North Dakota,
Wyoming, Montana, and Alberta.
Revised diapwsis. Vertebral colimm
with significantly fewer total ctMitra (65
mean) than the other species, with approxi-
mately twelve fewer trnnk vertebrae (25
mean) and eight fewer monospondylous
caudal centra ( 17 mean ) . Distance be-
tween anal fin insertion and the end of the
vertebral column relatively short, with
dorsal fin terminating close to caudal fin.
Caudal lepidotrichia 19-20 rather than 23-
27. Ascending processes of parasphenoid
perpendicular to the main anteroposterior
parasphenoid axis; more posterior place-
ment of parasphenoid tooth-patch. Pari-
etals squared in outline. Marginal teeth
simple pointed cones, palatal teeth usually
stout styliform crushers. Supraorbital sen-
sory canal not entering parietal. Excava-
tion of orbital notch in frontal relatively
larger. Dentary with additional horizontal
shelf of coronoid articulation surface adja-
cent to lingual border of alveolar ridge;
coronoid articulation surface extensive,
overlapping ventral half of ramus; dentary
with pronoimced arch rather than gradual
curve in ventral outline. Greatest known
standard-length 510 mm.
Introduction
Jordan (1927) described Kindleia fra^osa
as a new genus of cichlid fish from the Late
Cretaceous Edmonton Formation of Al-
berta. This tentative placement of Kindleia
within the Cichlidae was largely the result
of his misinteipreting the splenial tooth
plates for fused lower pharyngeal bones
(Estes, 1964). One month later, Russell
( 1928a) independently published a descrip-
tion of Stylomyleodon lacus, a new fossil
amiid from the Late Paleocene Paskapoo
Formation of Alberta, and referred other
specimens from the Edmonton Formation
of Alberta to the same species. His descrip-
tion also included a dentary and palatal
teeth modified tor crushing. His relegation
of the genus to the Amiidae was based on
a correct inteipretation of the "splenial"
(= coronoid) tooth plates (Estes, 1964).
He suggested a relationship of Stylonujle-
odon to Platacodon nanus ( at that time
erroneously considered an amiid; see Estes,
1964) with the essential difference being
hemispherical rather than Hattened tooth
crowns.
Jordan later ( 1928 ) noted the similarity
of the two genera Kindleia and Stylomyle-
odon and asserted the prior claim of his
name Kindleia. Although he made no com-
ment on Russell's attributing Stylomyleodon
to the Amiidae, he rejected Russell's com-
parison of that genus with Platacodon on
the basis of Marsh's earlier conviction that
the latter was mammalian. In reply to
Jordan, Russell ( 1928b ) defended the valid-
ity of his genus on the supposition that its
dentary was distinct from that of Kindleia,
although he did agree on the similarity of
teeth and jaw fragments of the two genera.
Russell (1929) further attempted to vali-
date Stylomijleodon as a genus by com-
paring his type with new specimens
collected by Princeton University. This new
material confirmed his association of the
maxilla-dentary and palatine-coronoid den-
titions, and also substantiated his interpreta-
tion of Stylomyleodon as an amiid in which
the coronoid teeth were specialized for
crushing. He also admitted that there was
insufficient Platacodon material to deter-
mine any conclusive similariti(\s with Stylo-
myleodon, but, r(>f(Mring to Hatcher's (1900,
1901 ) work, did insist that Platacodon was
a fish. Simpson (1937) reported finding
additional specimens of Stylomyleodon Rus-
sell in the Fort Union Formation at Crazy
Mountain Field sites of Montana.
Estes ( 1964 ) , from his studies of amiid
material from the Lance Formation of Wyo-
ming, observed that whereas the type
dentary referred by Russell to Stylomyleo-
40
Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
don was the pcsterior portion of an amiid
dentary, Jordon's type was the anterior
portion. From this fact he confirmed the
synonymy of Stylomtjleodon with the genus
Kindleia, at that time beUeving that it was
generically separated from Amia. Janot
( 1967 ) agreed with Estes on the synonymy
of Stylomijleodon with Kindleia, but did not
find sufficient cause to distinguish Kindleia
generically from A7?j/fl. Russell (1967) con-
tinued to leave the nomenclatural problem
of Stylomyleodon-Kindleia unsettled. Estes
and Berberian ( 1969) studied material from
the Late Cretaceous Hell Creek Formation
of Montana and confirmed Janot's proposi-
tion that Kindleia is a synonym of Amia.
They also suggested the possibility of
synonymy of A. fra<i,osa with A. keJireri
(Middle Eocene, Germany), A. russelli
(Late Paleocene, France), A. munieri
(Early Oligocene, France), and Paromiatus
gurleyi (Early Eocene, Wyoming), but
postponed formal synonymy of A. jrag,osa
with the prior name A. kehreri (Andreae,
1892), pending more detailed study of
Early and Middle Cenozoic specimens
from Europe. Estes and Berberian (1969:
10) concluded that the minor variations
that separated A. fra<^osa and its related
forms in Europe from A. calva are "super-
ficial and essentially primitive," and indi-
cated that the group might be close to the
ancestry of the Recent species A. calva.
A nearly complete skull from the Eocene
Willwood Formation of Wyoming (Fig.
16), two axial skeletons from the Eocene
Green River Formation of Wyoming ( Plate
1 ) , and a sample of disarticulated elements
from the Late Cretaceous, Paleocene, and
Early Eocene have yielded more informa-
tion on the osteology of Amia fragosa. Estes
(1964), O'Brien (1969), and Estes and
Berberian (1969) have studied this species
in detail, and I therefore discuss these speci-
mens only as they modify conclusions
reached by those studies.
Fossil Record
In addition to the stratigraphic list given
by Estes and Berberian (1969: 14, table 1)
of major freshwater deposits carrying A.
fragosa, three new localities are recorded
here: the Late Paleocene Silver Coulee
local fauna of the Fort Union Formation,
Wyoming, and the Early Eocene Willwood
and Wind River formations, Wyoming. The
luajor deposits in which remains of A.
fragosa have been found are summarized in
Table 18. Estes and Berberian (1969: 10)
state that the stratigraphic range of A.
fragosa extends from the Late Cretaceous
through at least Middle Eocene time in
North America. The earliest deposit in
which remains of A. fragosa have been
found is the Late Cretaceous (Campanian)
Oldman Formation of Alberta, and the
latest deposit is the Middle Eocene (Brid-
gerian) Bridger Formation of Wyoming.
Cavender (1968: 128), however, de-
scribes Amia scales from the Late Eocene
( Duchesnean ) Clarno Formation of Ore-
gon. Although these small scales (approxi-
mately 2 mm in length) are not as robust
as those of A. fragosa, they are more ossified
than A. scutata and A. calva scales. These
scales, along with the scales from the Horse-
fly River Beds of British Columbia (UMMP
collections) cannot at present be identified
as to species. They are best referable to
Amia sp., since no identifiable A. fragosa
elements have been found later than
Bridgerian and no specific character-states
for scales of A7nia have yet been deter-
mined.
Description
Neurocranium. Estes (1964: 29) stated
that the greater length of the basioccipital
and the presence of a second pair of aortal
supports in Amia calva indicated that the
basioccipital posterior to the spinal (inter-
vertebral) arterial foramina included only
one fused vertebra in Amia fragosa instead
of the two found in A. calva. O'Brien
(1969: 42) observed a similar condition in
two complete A. fragosa specimens from
the Edmonton Formation of Alberta. Estes
and Berberian (1969: 2-3) found nine
basioccipitals with one fused vertebra and
eleven with two fused vertebrae from the
V
Fossil Amiids • Boreske 41
Fig. 16. Amia fragosa MCZ 9264, Early Eocene, Willwood Formation, Wyoming; A, dorsal; B, ventrol.
Hell Creek Formation of Montana and in-
terpreted this as a variation in A. frci'^osa
not observed in the Lance sample. Janot
( 1967 ) noted that basioccipitals of the
European Late Paleocene Amia sp. also
showed this variation. Estes and Berberian
(1969: 2) suggested that a weak tendency
for fusion of vertebrae could be correlated
with increasing size, and that such varia-
tion might possibly e.xist in the Recent
species as well. Fifty Recent A. calvo skele-
tons examined, with a size range of 100-480
42 Bullctiti Museum of Comparative Zoology, Vol. 146, No. 1
mm SL, have the first two vertebrae fused
to the basioceipital. Three articulated and
twenty-two disarticulated Eocene and
Oligocene Amia uintaensis Ijasioccipitals
all have two vertebrae fused to the basioc-
eipital. Unfortunately, in specimens of A.
fni'^osa (MCZ 9264, PU 13261) having a
visible parasphenoid, the basioceipital re-
gions are poorly preserved. There is a pos-
sibility that the Lance sample by chance
contained only specimens with one fused
vertebra since there are only six specimens
known. Until more specimens of A. jra<iosa
and A. uintaensis with intact basioccipitals
become available, it is difficult to discuss
this point further.
Fig. 17. Comparison of parasphenoids of Amio spp.:
c, Amia calva. Recent, after Janot, 1967 (c'' := dorsal, c^'
= ventral); f, A. fragosa (ventral); u, A. uinfaensis
(ventral).
The length of the A. fra<j,osa parasphe-
noid posterior to the ascending processes is
10 percent shorter and slightly wider than
in A. calva, with the ascending processes
more posterior than in the Recent species
(Fig. 17). The proportion of the length
posterior to the processes to the length
anterior to these processes (0.780) is not
as small as in A. uintaensis (0.704) or as
great as in A. calva (0.900), and, on the
basis of this small sample, A fragosa is inter-
mediate among the three species for this
character. The region posterior to the
processes also appears more convex than in
A. calva, but not as convex as in A. uijitaen-
sis. The ascending processes are almost
perpendicular to the main anteroposterior
axis of the parasphenoid. Those of A.
fragosa form an approximately 85-degree
angle with the parasphenoid axis, while the
ascending processes of both A. calva and
A. uintaensis form approximately 70-75-
degree angles. The mid-ventral surface of
the parasphenoid bears small, sharp, con-
ical teeth. This tooth-bearing surface of A.
fragosa terminates anteriorly toward the
middle of the ascending processes, whereas
in A. calva this region narrows to a point
and extends to the posterior end of the
vomers (Fig. 17). In A. uintaensis, this
region also extends to the vomers, but
covers a wider surface area in the anterior
region than in A. calva. Nearly all the
tooth-bearing surface of A. frasj^osa lies in
the posterior half of the parasphenoid,
while in A. calva this surface is centered
between the posterior and anterior areas;
in A. uintaensis two-thirds of this surface
lie in the anterior region of the para-
sphenoid. The entire tooth-bearing surface
of A. fragosa is wider than that of A. calva,
since the anterior half of the tooth-bearing
surface tapers anteriorly in A. calva, while
in A. fragosa the anterior portion maintains
a more constant width. The basic outline of
the tooth-bearing surface in A. fragosa is
subrectangular; that of A. calva is more
tear-drop shaped, with the anterior apex
widened and extended to the vomers. The
two posterior parasphenoid flanges are
Fossil Amiids • Boreske 43
more splayed in A. jra<^osa tlian in A. caJva
or A. uintacnsis, and overlie three-fourths
of the basioecipital length. As Estes ( 1964:
29) notes, there is a relatively greater dor-
soventral parasphenoid thiekness in A.
fniiiosa than in A. calva. The parasphenoid
of A. iiiiitoensis is proportionately more
massive than that of A. frai!,osa: this mas-
siveness, however, is probably a function
of its greater size.
In A. fraiiosa, as in A. uintaensis, the
extrascapular is tear-drop shaped, being
narrow at the midline and expanded dis-
tally, while in A. calva and A. sciitata, it is
more strap-shaped and longer at the mid-
line. The proximal anterior corner is
squared off, as in A. scutata and A. calva.
The anterior edge is distally concave at
the pterotic-extrascapular suture, and the
posterior edge is convex, particularly
toward the distal end, which is straight
rather than curved as in the other species
of Amia. A. fragosa and A. uintaensis lack
the posterolateral projection displayed in
A. calva.
The suprascapular resembles that of A.
calva, except that the distal edge is rela-
tively straight, rather than incurved. The
posterior border is also straight, while in
A. calva there is generally a slight concavity
in the middle of this edge; in A. uintaensis
this border is convex.
The pterotic extends further anteriorly
than in A. calva, but not to the extent that
it does in A. uintaensis or A. scutata. The
dermosphenotic-pterotic suture is directed
posterolaterally in A. fragosa and antero-
laterally in A. uintaensis, A. scutata, and
A. calva. As in A. uintaensis and A. scutata,
this bone in A. fragosa is narrower an-
teriorly than posteriorly, whereas in A.
calva the widths of these ends are relatively
equal.
The dermosphenotic in A. fragosa is about
the same relative size as in the other species
of Amia. The anterior angle that forms the
posterior border of the orbit is slightly more
pronounced than in A. scutata and A. uin-
taensis, and considerably more so than in
A. calva (Fig. 28).
The parietal in A. fragosa is character-
istically square, whereas in A. calva, A.
scutata, and A. uintaensis it is longer than
wide. The length of the parietal relative
to that of the frontal is less than in A. calva
and A. scutata and about the same as in
A. uintaensis. The characteristic deep ex-
cavation in the frontal for the orbit is
displayed in all available specimens of
A. fragosa. This led Estes (1964: 36) to
postulate the presence of supraorbital bones,
but the articulated specimens figured by
O'Brien (1969) show that this was not the
case. As Figure 28 shows, the ratio of
orbital depth to length is greater in A.
fragosa than in tlie other Amia species. As
noted in the preceding section on the cranial
morphometries of the Recent A. calva, it is
difficult to assign a specific character-state
of parietal/frontal proportions to any of the
individual fossil Amia species because of
the similarity in parietal /frontal propor-
tions (Table 7). It is apparent, however,
that the frontals of the earlier species A.
fragosa and A. uintaensis are longer relative
to parietal-length than in the mid-Tertiary
A. scutata or Recent A. calva. This feature
is useful in comparing A. fragosa with these
two species, but ineffective in distinguishing
it from A. uintaensis.
As Estes and Berberian (1969) noted,
the nasal displays a bifurcation of the an-
terior border that is lacking in A. calva.
The bifurcation is also present in A. uintaen-
sis, and the bone has approximately the
same outline and size relative to head size
as the other forms. All available specimens
of A. fragosa show that the nasals lie much
closer to the frontals than in A. calva, A.
scutata, or A. uintaensis. Although Estes
( 1964 ) states that the lacrimal conforms
closely with that of A. calva, his restoration
lacks the small posterior notch in A. fragosa
which accommodates the anterior process
of infraorbital 2. The lacrimal in A. fragosa
is evenly tapered at the posterior end, and
is anteroposteriorly longer than in other
Amia. It is also more dorsoventrally convex
than in A. scutata and A. calva.
As in A. scutata, infraorbital 4 in A.
44 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
fragosa is much more dorsoventrally ex-
panded than in A. calva, with the antero-
posterior length extending almost to the
anterior edge of the preoperculum. Infra-
orbital 4 of A. fragosa and A. scutata is
more concave at the dorsal edge, and much
more convex ventrally. The pit-line marks
extend further ventrad than is indicated in
the reconstruction by Estes ( 1964 ) .
Branchiocranium. In A. fragosa, the
supramaxilla is relatively shorter than in
A. iiintaensis, A. scutata, and A. calva, with
a greater curve in the maxillo-supramaxil-
lary suture. The dorsoposterior corner in
A. fragosa tends to be angular, as in A.
scutata, whereas in A. calva and A. uintaen-
sis it is more rounded. The supramaxilla is
deeper and more truncated at the anterior
end than in other species of Ainia.
There is a dorsal shelf adjacent to the
lingual border of the alveolar ridge which
widens the anterodorsal surface of the den-
A. fragosa A. uintaensis A. scutata A. calva
A. fragosa
A. uintaensis
A. scutata
Fig. 18. Comparison of mandibles of
(transverse sections and ventral views).
spp.
tary (Fig. 18). This shelf is lacking in
A. calva and A. uintaensis, in which the
coronoid articulation surface slopes directly
downward from the alveolar ridge. This
region of the lingual dentary surface under-
lying the coronoids extends more ventrad
at the symphyseal edge than in A. calva,
and distinctly overlaps the ventral part of
the ramus. There is no such overlapping in
A. calva; the dorsal and venti'al halves of
this region separate to form Meckel's groove.
The anterodorsal section of the dentary in
A. uintaensis overlaps the ventral half, but
not to the extent that it does in A. fragosa,
and as the coronoid articulation surface is
thicker, this thickened area of bone forms
the dorsal wall of Meckel's groove as in
A. calva (Fig. 18). As Estes (1964: 36)
noted, the coronoid teeth are styliform and
extend almost to the ventral border of the
ramus at the anterior end; in contrast, the
coronoid teeth of A. calva, A. scutata, and
A. uintaensis are pointed and the coronoids
do not extend as far ventrally as in A.
fragosa. The anterior half of the dentary
length is more curved than in A. calva, A.
scutata, and A. uintaensis (Fig. 18). This
is displayed in the MCZ 9264 specimen
( Fig. 16 ) , in which this curve approximates
a 120-degree angle at the midpoint of the
alveolar ridge. The outline of the dentary
differs from that of A. calva and A. uintaen-
sis in that the anterior end maintains an
almost constant width up to the sharp
curve at the midpoint of the alveolar ridge,
at which point it widens noticeably. When
the outline and curvature of the anterior
end of the dentary of A. fragosa are com-
pared with those of other species, the result-
ing difference appears to be correlated with
A. fragosa's relatively smaller mandible/
head ratio (Table 7), smaller mouth gape,
and its wider cranial roof (Fig. 15).
Post-cranial Skeleton. On the basis of
specimens having only the lateral surface of
the vertebral column exposed, it was con-
cluded that centra of A. fragosa are indis-
tinguishable from those of A. scutata and
A. calva. Small disarticulated vertebrae are
also basically similar in morphology, there-
Fossil Amiids • Boreske 45
fore it is impossible to differentiate A.
frau^osa, A. sciitata, and A. calva. The mid-
trnnk vertel^rae of A. fra^osa, A. sciitata,
and A. calva differ from A. uintaensis mid-
trunk vertebrae, which are generally larger,
and snbtriangular rather than ovoid. A.
fragosa does, however, have a vertebral
column that differs proportionately and
meristically from that of other species ( Fig.
31). It has a significantly smaller total
number of centra than the other species,
with approximately 12 fewer trunk verte-
brae and 8 fewer monospondylous caudal
centra. It has the same number of diplo-
spondylous caudal centra as A. calva and
A. scutata; the number of fused hypurals is
also generally the same (Table 9). The
low number of total vertebrae in A. fragosa
is reflected by its shorter, deeper-bodied
shape. The distance between the anal fin
insertion and the end of the vertebral
column is relatively shorter than in the
other species. The dorsal fin also terminates
closer to the caudal fin than in any of the
other species of Amia (Plate 1; Estes and
Berberian, 1969: 10). A. fragosa has fewer
caudal lepidotrichia (19-20) than the other
species of Amia (23-27). The head/
standard-length ratio of A. fragosa is greater
than that of A. calva, but is not significantly
different from that of A. uintaensis or A.
scutata (Table 3). The latter case is true
despite the greater number of vertebral
centra in A. uintaensis and A. scutata; this
disparity may be explained largely by the
fact that the A. fragosa skull itself is rela-
tively shorter than that of the other two
forms, particularly A. uintaensis, which has
a greater head/ standard-length ratio than
A. fragosa. Thus head/ standard-length does
not significantly reflect the length of the
vertebral column, but may be used as a
character with this qualification in mind.
The known total-length of A. fragosa falls
within the range of A. calva and below that
of A. uintaensis (Tables 1-2).
Discussion
Marsh (1871: 105) described Amia new-
berrianus and Amia depressus on the basis
of disarticulated vertebrae and cranial ele-
ments from the Bridger Formation of Wyo-
ming. His main criteria for distinguishing
these forms from A. calva and from each
other were that the chordal foramen of A.
newherrianus was "considerably above the
center in the dorsal vertebrae," and that
A. depressus possessed broader vertebrae
than A. newherrianus and lacked the me-
dian groove on the lower surface of the
centra. The vertebrae indicated that both
species were approximately the size of
A. calva. Osborn et al. (1878: 102) noted
that since Marsh gave no measurements,
"the reference to Amia depressus cannot be
certain." Marsh further noted that these
specimens belonged to the Yale College
Museum, but the specimens now seem to
have been lost. Marsh had apparently as-
sumed that the characteristics of one verte-
bra represented those of the entire vertebral
column and was unaware of intracolumnar
variation in height/ width proportions, aortal
facet morphology, and position of chordal
foramen in the vertebral coliunn of Amia.
I infer from Marsh's report that the type
specimen of A. depressus is probably a first
to third trunk vertebra, since the aortal
grooves are lacking (Fig. 11) and vertebral
width exceeds height (Fig. 14). Using the
position of chordal foramen as a character
distinguishing A. neivberrianus is undiag-
nostic since the position of the chordal
foramen changes in relation to the relative
position of the vertebra along the column
(Fig. 12). Therefore, on the basis of
Marsh's undiagnostic characters and the
similarity in size and morphology of the
vertebrae to those of A. fragosa and A.
calva, I consider both A. depressus and A.
newherrianus as nomina duhia.
Leidy (1873a: 98) descrilied Amia graci-
lis from a single trunk vertebra, also from
the Bridger Formation of Wyoming. He
noted that the centrum has two "oblong
fossae" ( aortal facets ) instead of the charac-
teristic pair of v(>ntral ridges found in Amia
calva. The size of the centrum indicated to
Leidy that A. gracilis was a smaller species
than A. calva (Leidy, 1873b). The vertebra
46 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
(ANSP 5360) corresponds approximately
to the twelfth trunk vertebra in Ainia, since
the aortal facets are oblong and unridged
(Fig. 11). Although A. gracilis is small, it
falls well within the size range of A. fra^osa
and A. calva, and is considered as a nomen
duhium.
Estes and Berberian (1969: 10) suggested
the possibility of synonymy of Paramiatus
gurleyi (Plate 1) and Atnia fra<i,osa with
the European Amia kehreri (Plate 2) on
the basis of the close proximity of dorsal
and caudal fins for the former and similar-
ity of skull elements and teeth for the latter.
It was shown in the previous section that
Paramiatus <i,urleyi conforms not only to
A. kehreri, but also to North American spec-
imens of A. frafi,osa on the basis of body
morphometries and meristics. Cranial mor-
phometries were also shown to be similar.
In addition, an X-ray (FMNH X2201) of
the Paramiatus <^urleyi skull reveals infra-
orbitals 4 and 5 to be longer than Romer
and Fryxell (1928) and Estes (1964) had
noted. The two infraorbitals extend pos-
teriorly to the anterior edge of the pre-
operculum as they do in A. frau^osa (UA
5398) from the Late Cretaceous Edmonton
Formation of Alberta. The X-ray has also
revealed a displaced left vomer with 26
styliform teeth which has been rotated
through the skull roof and covered with
matrix. All other cranial elements conform
morphologically with other Cretaceous and
Eocene specimens of A. frafiosa. These
additional similarities of Paramiatus gurleyi
and A. fragosa confirm the synonymy of
these two species which was suggested by
Estes and Berberian (1969: 10), and I
therefore include Paratniatus gurleyi in the
synonymy of Amia fragosa.
Comments on Related European Forms
Amia kehreri was described by Andreae
(1892, 1895) from a caudal region, infra-
orbital 4, disarticulated trunk vertebrae,
and a left operculum from Middle Eocene
(Lutetian) deposits at Messel bei Darm-
stadt (specimens at the University of
Heidelberg, Andreae Collection). On the
basis of specimens from the same deposit
(BMNH P33480, Plate 2; P33488), it con-
forms morphometrically with A. fragosa in
head/ standard-length, pectoral fin insertion/
standard-length, mandible/ head-length, and
operculum-length/operculum-depth ( Ta-
bles 3, 7). The distance between the anal
fin and the end of the vertebral column
exceeds that of the North American A.
fragosa specimens, but is less than in A.
scutata or A. calva. The parietal/ frontal
ratio is also greater than in A. fragosa, but
smaller than in A. scutata or A. calva.
Meristics of vertebral elements as well as
the cranial characters discussed by Estes
and Berberian ( 1969 ) also conform with
those of A. fragosa. I agree with Estes and
Berberian ( 1969 : 10 ) that only differences
in temporal and geographical factors appear
to distinguish Amia kehreri from A. fragosa;
any osteological dissimilarities are of minor
significance.
The Middle Eocene European Geiseltal
deposits contain numerous amiid fossils;
according to Estes and Berberian ( 1969 )
some showed resemblances to A. kehreri.
This material is currently being described
by Anna Jerzmanska, Uniwersytet Wroclaw-
ski, Wroclaw, Poland.
Another related form is Amia valencien-
nesi from the Eocene of Puy-de-D6me,
France. Agassiz ( 1843 ) described the form
from one complete specimen and an an-
terior region of another (BMNH P446,
27736). Piton (1940) reviewed these .speci-
mens along with new material collected at
the same locality. A. valenciennesi also re-
sembles A. kehreri in its vertebral number
of 68 centra, close approximation of dorsal
and caudal fins, and an infraorbital 4 larger
than infraorbital 5; these similarities indi-
cate that synonymy with A. kehreri is in
order. The name A. valenciennesi precedes
A. kehreri, and thus has priority.
Estes and Berberian (1969: 7) showed
that Amia russelli Janot ( 1966, 1967 ) from
the Late Paleocene of France resembles A.
fragosa in (!) square parietals, (2) similar
Fossil Amiids • Boreske 47
parietal/frontal ratio, (3) largo orbital ex-
cavation in frontal, and (4) similar opc>r-
cnlnni height/width ratio. Thns A. russclli
conforms with many of the most distinct
characters of A. kehreri and A. valencien-
nesi, and should be considered a synonym
of the latter.
Estes (1964) re-evaluated Dechaseaux's
( 1937) redescription of the Early Oligocene
Amia munieri from France and noted simi-
larities with A. fraiS,osa which included ( 1 )
styliform vomerine teeth, (2) branchiostegal
rays rounded distally, ( 3 ) larger infraorbital
4 than infraorbital 5, and (4) similar
parietal /frontal proportions. The principal
difference between the forms is the small
excavation for orbits in A. munieri. Since
Dechaseaux's and Estes' studies, the speci-
men (MNHN R4632, skull and associated
cranial and postcranial elements) is being
further prepared to display the cranial roof
and palate more extensively. The frontal
lacks a prominent excavation for the orbits
as Estes (1964: 40) has noted, and in this
feature A. munieri resembles A. sctitata and
A. calva. A. mimieri is a very important
form because it represents the only com-
plete amiid specimen known from the Early
Oligocene, and, as noted, it displays inter-
mediate morphology of the cranial features
among the species of Amia. A. munieri
occurs very late in time in relation to the
last known occurrence of A. jragosa in
North America, and because there are no
complete specimens known from this age,
it represents a stage of evolution among the
amiids that is not found in North America.
Lehman ( 1951 ) described Fseudamia
lieintzi (Troms0 Museum Naturhistorisk
collections, Troms0, Norway) from a fairly
complete articulated sptx-imen and two
skulls from probable Eocene deposits in
Spitzbergen. He differentiated this form
from Amia on the basis of ( 1 ) Sinamia-\\ke
metapterygoid and (2) presence of a con-
cave notch on the dorsoposterior border of
the operculum. Estes ( 1964 ) noted that
Lehman was incorrect in his interpretation
of tlie nature of the metapterygoid and
operculum, and therefore suggested that
Fseudamia might be placed in the genus
Amia. From the examination of Lehman's
plates, it appears that this form resembles
A. fra^osa in its deep-bodied shape and
low parietal /frontal ratio (approximately
0.410), and that it may be synonymous with
A. valenciennesi and A. kehreri. I'urther
preparation would possibly be helpful in
uncovering palatal teeth, whose moiphology
would aid in a more definitive description.
Although the exact age of the Eocene
deposit in which the specimen occurred
is uncertain, this Spitzbergcni locality, if
Early Eocene, lies on the possible migration
route of amiids (and other vertebrates)
between North America and Europe.
Amia uintaensis (Leidy, 1873)
Protamia tiintacnsis Leidy, 1873a: 98.
Protamia media Leidy, 1873a: 98.
Pappichthy.s plicatus Cope, 1873: 635.
Pappichthtjs sclerops Cope, 1873: 635.
Pappichthy.s laevis Cope, 1873: 636.
Pappichthijs symphysis Cope, 1873: 636.
Pappichthys corsonii Cope, 1873: 636.
Pappichthys meditis Cope, 1884: pi. 4.
Amia ivhiteavcsiana Cope, 1891: 2.
Amia macrospondyla Cope, 1891: 2.
Holotype. ANSP 5558, anterior tiimk
vertebra.
Paratypes. ANSP 8044, first anterior
trunk vertebra; ANSP 3151, three posterior
trimk vertebrae; ANSP 5622, basioccipital.
Type locality and horizon. Henrv's Fork.
North half of section 5, T 12 N, R 111 W,
Sweetwater County, Wyoming; Bridger
Formation.
Age rouge. Torrejonian (Middle Paleo-
cene) to Chadronian (Early Oligocene).
Hypodi^m. Paleocene. Fort Union For-
mation, Wyoming and Montana: PU 17117,
maxillary; PU 17068, vertebrae and denta-
ries; PU 162.36, disarticulated skull and
trunk vertebrae; CM 25364, dentary; PU
17064, trunk vertebrae. Tongue River For-
mation, Montana: PU 20578, basioccipital
and vertebrae. Paskapoo Formation, Al-
berta: ROM 4653, vertebrae.
Eocene. Will wood Formation, Wyo-
ming: PU 21173, basioccipitals; PU 17227,
basioccipital and trunk vertebrae; PU 17649,
1
48 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
portion of cranium; PU 18760, skull frag-
ments, dentary, and vertebrae. Wasatch
Formation, Wyoming: AMNH 4635, dentaiy
and maxilla. Golden Valley Formation,
North Dakota: PU 18568, basioccipital.
Green River Formation, Wyoming: USNM
18147, skull fragments and vertebrae;
AMNH 785, complete caudal region; PU
13865, nearly complete specimen; MCZ
12916, disarticulated skull and associated
vertebrae. Wind River Formation, Wyo-
ming: AMNH 2437, dentary and skull frag-
ments. Bridger Formation, Wyoming: CM
25362, portion of cranium and vertebral
column; AMNH 4631, portion of cranium
with dentaries, gular, and basioccipital;
USNM 170976, maxilla; YPM 6238-6240,
6242, 6244, 6250-6253, 6257-6258, vertebrae
and basioccipitals; USNM 170973, 5450,
3962, 3963, 3966, PU 20523, 10101, ANSP
2337-2339, vertebrae; USNM 2181, ANSP
5632, trunk vertebrae; USNM 3959, trunk
and caudal vertebrae; ANSP 5580, mid-
trunk vertebra; AMNH 2539, anterior por-
tion of a left dentary, two premaxillae, right
quadrate, left epihyal, anterior portion of an
ectopterygoid, three trunk vertebrae, and
numerous fragments of angular; USNM
3965, left dentary; USNM 3968, anterior
dentary fragment; AMNH 2570, pre-maxil-
lary fragment, fragments of angular, left
quadrate fragment, trunk vertebra frag-
ment, and a caudal vertebra; USNM 3960,
PU 10099, 10110, vertebrae and a ural cen-
trum; USNM 5476, basioccipital; USNM
3961, left dentary fragment. Washakie For-
mation, Wyoming: FMNH 27465, 4509, ver-
tebrae. Uinta Formation, Utah: CM 2382,
maxillary fragment.
Oligocene. Cypress Hills Formation,
Saskatchewan: NMC 6197, trunk vertebra;
NMC 6198, caudal vertebra.
Known distribution. Montana, Wyo-
ming, Utah, North Dakota, Alberta, and
Saskatchewan.
Revised diagnosis. Vertebral column
with approximately 20 more vertebral seg-
ments in total number (85) than A. fragosa,
and five fewer trunk centra (31) and five
more diplospondylous caudal vertebrae
(21) than in the other long-bodied forms,
A. scutata and A. calvo. Mid-trunk verte-
brae subtriangular rather than ovoid. Pa-
latal teeth sharp, greatly curved inwardly.
Between 40-45 vomerine teeth as compared
with 15-17 in A. fragosa, A. scutata, and
A. calva. Hyomandibular more deeply
notched between opercular process and
extensor (dorsal) surface than in other
species; opercular process relatively larger.
Angle between alveolar ridge and exterior
surface of the dentary forms a more acute
angle than in the other species. Mandibular
ramus less curved than in other species, so
that angle between symphyseal ends of
dentaries is relatively narrow. Greater
mandible/head ratio (0.693) and head/
standard-length ratio (0.322) than any of
the other forms: A. uintaensis has a head
relatively longer and a mouth gape rela-
tively wider than do other species. Most
specimens are significantly larger than the
other species, with a relatively greater de-
gree of ossification of all bones. Greatest
known standard-length 800 mm.
Introduction
Leidy (1873a) reported numerous dis-
articulated vertebrae of a fossil fish related
to Amia from the Bridger Formation of
Wyoming. He distinguished a new genus
Protamia from Amia by its "two oval fossae"
( aortal facets ) on the ventral surface of the
centrum, and by large vertebrae character-
istically with a much greater width to
height proportion. Hijpamia, another new
genus from the same locality which Leidy
also related to Amia, was characterized by
also being larger than A. calva, and by
vertebrae whose sides converged into a
"medium prominence excavated into a pair
of oval fossae" deeper than those of Pro-
tamia. Later ( 1873b ) , Leidy published a
more complete and illustrated account of
the various species of the new genera
Protamia and Hypamia. In the same year
Cope ( 1873 ) described a new amiid genus,
also from the Bridger Formation, which he
named Pappichthys. He distinguished this
new genus from Amia by the "presence of
Fossil Amiids • Boreske
49
only one series of teeth, instead of several,
on the bones about the mouth." Osborn
et al. ( 1878 ) reported other finds of Pci})-
picJitJiys from the Bridger Formation which
seemed to fit Cope's description. Cope
(1884) further discussed his new genus,
and rejected Leidy's prior nomenclatiue
and description.
New ton ( 1899 ) discussed this nomencla-
tural controversy and asserted the validity
of Leidy's genius Protamia, since Cope's
later diagnosis \\'as no more effective in
characterizing the new genus than Leidy's
prior one. Newton bc^lieved that Cope's
description of PappicJitlitjs as having only
a single row of marginal teeth was taxo-
nomically undiagnostic, since this condition
would also include A. calva. Romer and
Fryxell ( 1928 ) accepted Leidy's earlier
description and genus as diagnostic, and
referred PappiclitJujs to Protamia. They
also mentioned Hypamia but found little
to distinguish it from Amia.
Hussakof (1932) continued to use Cope's
name, however, and reported large speci-
mens of Pappichthys from the Eocene of
Mongolia. He also noted Cope's error in
diagnosing the tooth characteristics of the
genus, since Pappichthys {Protamia) has
several rows of small teeth on the "splenial
bone." In comparison with Amia he noted
"points of difference in nearly every bone
available for comparison," and concluded
that Pappichthys was a valid genus, "not
merely a group of large-sized extinct species
of Amiatus."
Estes (1964), like Romer and Fryxell
(1928), referred Pappichthys to Protamia,
and reported several vertebrae and a maxil-
lary fragment from the Cretaceous Lance
Formation of Wyoming. He inteipreted the
increase in breadth over thickness of the
vertebrae as a po.ssible "function of in-
creased size," a condition that would also
allow for tlie comparatively more massive
nature of the maxillary fragment. He also
considered the retention of this genus as
arbitrary until enough materials were avail-
able. Janot (1967) did not consider this
single distinguishing characteristic of the
vertebrae as sufficient foundation for the
erecti(m of a new genus, and therefore sug-
gested relerring Protamia to Amia. Estes
et al. (1969) concurred with Janot in
synonymizing Protamia with Amia. The
present study confirms tliis synonymy;
Leidy's species (1873a) has priority and
the valid name of this fish is thus the oldest
specific name, Amia uintacnsis.
Revision of all forms referred now or in
the past to Protamia is much needed, for
these large amiids were diagnosed on char-
acters of isohited vertebrae and skull frag-
ments. This study gives more useful
diagnostic characters that provide a basis
on which the taxonomy of this group can be
established.
Fossil Record
The major deposits carrying remains of
Amia uintacnsis (Table 18) range in age
from Middle Paleocene to Early Oligocene.
Middle Paleocene specimens occur in the
Fort Union, Tongue River, and Paskapoo
formations and consist mostly of isolated
and broken centra, and dentary and maxil-
lary fragments. A nearly complete skull
(PU 162.36) with associated trunk and
caudal centra from the Bear Creek local
fauna of Montana (Fort Union Formation)
is the only articulated specimen from the
Late Paleocene. The Eocene material in-
cludes one complete articulated specimen
(PU 13865), one complete caudal region
(AMNH 785), and a disarticulated skull
(MCZ 12916) from the Creen River Forma-
tion. PU 13865 (Plate 3) has the axial
skeleton intact in matrix, with a dislocated
fifth centrum that is the only one available
for three-dimensional measurements. This
is also the only specimen in which a com-
plete vertebral count can be taken. AMNH
785 provides excellent meristic information
for the caudal region (Fig. 8C). CM 25362,
from the Bridger Formation, consists of a
left palatal and opercular series and an
almost complete, disarticulated vertebral
column that permitted the taking of a series
of centrum measurements. Other skull frag-
ments and vertebrae occur in many deposits
50 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
throughout the Eocene (Table 18). The
latest occurrence of A. uintaensis is repre-
sented by two isolated centra from the
Cypress Hills Formation (Oligocene, Chad-
ronian ) .
Description
Neurocranhan. Posterior to the spinal
arterial foramina the basioccipital includes
two fused vertebrae. As the basioccipitals
display great variation in the morphology of
the articular surface, it is difficult to char-
acterize this form on the basis of this
feature. However, the articular surface is
generally kidney-shaped, with dorsal in-
dentations bet\veen the neural facets, and
ventrally there is an indentation distal to
the aortal facets. In A. froii^osa and A.
calva the basioccipital has ovoid articular
surfaces with no dorsal indentations be-
tween the neural facets (Estes, 1964: 29,
fig. 15). In lateral view the distal articular
surface of the A. uintaensis basioccipital is
not perpendicular to the parasphenoid
flanges; the dorsal half of this surface is
more anteriorly directed than the ventral
half.
The parasphenoid is longer relative to
its width than it is in either A. calva or
A. fragosa, primarily in the region anterior
to the ascending processes. At the point
nearest the ascending processes, it lacks
the pronounced convexity and die accom-
panying anterior lateral notches found in
A. calva and A. fragosa. The ascending
processes are slightly less anteriorly oriented
in ventral view than in A. calva, but more
so than in A. fragosa ( Fig. 17 ) . The region
posterior to the ascending processes is rela-
tively shorter than in A. fragosa or A. calva;
it is also more massive and more ventrally
convex than in the other two forms. The
posterior parasphenoid flanges resemble
tliose of A. calva more tlian A. fragosa in
outline as well as juxtaposition; those of
A. fragosa are more laterally splayed than
in A. uintaensis or A. calva. The tooth-bear-
ing surface differs considerably from that of
A. fragosa and somewhat from A. calva in
outline and extent. As in A. calva, this
surface extends anteriorly to the vomers,
but its width is much greater and more
constant than in A. calva, which is narrowly
tapered anteriorly. Posteriorly, this surface
extends further than in A. calva, but not as
far as in A. fragosa. Approximately two-
thirds of the tooth-bearing surface lies
anterior to the ascending processes, while in
A. calva this area is anteroposteriorly cen-
tered, and in A. fragosa it is nearly all
posterior. The tooth-bearing surface covers
a greater portion of the ventral surface of
the parasphenoid than in A. fragosa or A.
calva; its basic outline is diamond-shaped,
with the anterior apex widened and ex-
tended to the vomers, while that of A.
fragosa is subrectangular and that of A.
calva is tear-drop shaped with the apex
sharply protracted anteriorly.
In A. uintaensis the distal edge of the
suprascapular is convex as in A. calva, while
in A. fragosa this edge is almost a straight
line. The posterior border is more rounded
distally than in A. calva and is convex rather
than concave.
In having the extrascapular rounded at
the distal border, A. uintaensis is the same
as A. calva and A. scutata, but differs from
both of them in that th(^ posterior border
is not concave, and from A. calva alone in
lacking the distal posterior process. The
anterior border is relatively straight, unlike
the condition in A. calva and A. scutata, in
which the lateral distal ends of the anterior
borders are directly posteriad. As in A.
fragosa the midline is shorter than in A.
scutata and A. calva.
As in A. scutata and in A. fragosa the
pterotic is narrower at the anterior than
posterior border, while in A. calva and, to
an extent, in A. scutata the ends are sub-
equal. As in A. fragosa they extend farther
anteriorly and adjoin the frontal s postero-
laterally. The dermosphenotic-pterotic su-
ture is anterolaterally directed, as in A.
scutata, but not as pronoimced as in A.
calva. The anterolateral edge of the pterotic
is indented and forms, witli the dermo-
sphcnotic, an additional concavity in the
outline of the cranial roof. Aside from this
Fossil Amiids • Boreske 51
anterior indentation, the lateral borders are
relatively straight, as eompared with the
smoothly coneave exterior sides of the
pteroties in A. scututa, A. calva, and A.
fruf!,osa. The posterior border forms a
smooth line, as in A. fragosa, and laeks the
small lappet that A. scutata and A. calva
display.
The dermosphenotie is similar to that ot
A. calva in relative size and outline, al-
though it does not jut as deeply into the
frontals. Its anterior border is rounded, as
in A. calva, rather than sharply angular, as
in A. fra^osa. The posterior half of the
outer lateral border is indented to form a
coneavity with the anterior tip of the
pteroties. The parietal in A. uintaensis is
elongated anteriorly, as in A. calva and A.
scutata, while that of A. jra^osa is relatively
square. The orbital excavation in the lateral
sides of the frontal is shallow as in A. calva
and A. scutata, while that of A. fragosa is
characteristically deep (Fig. 28). The sen-
sory canal cannot be determined. The
frontals are more elongated relative to
parietal length tlian in A. calva and A.
scutata; the parietal /frontal ratio is only
slightly smaller than that of A. fragosa
(Table 7). The distal lateral border tapers
anteromedially, and the anterior ends are
relatively pointed anteriorly, forming a deep
notch on the midline suture.
There is a slight bifurcation of the
anterior border of the nasal as in A. fragosa.
The nasal bones are relatively narrower
than in A. fragosa or A. calva, but are
otherwise similar in shape and relative size.
They are fairly well separated from the
frontals, as in A. calva and A. scutata, rather
than abutting them as in A. fragosa.
The lacrimal in A. uintaensis resembles
that of A. fragosa in general morphology,
although it lacks the posterior notch for the
anterior end of infraorbital 2 which is
present in the other species of Amia. The
lacrimal, like that in A. fragosa, is relatively
longer and more tapered posteriorly than in
A. scutata and A. calva. It is more dorsally
convex than in the other forms, but only
slightly more so than in A. fragosa.
The infraorbital 5 in A. uintai'nsis is
similar to that in A. fragosa and A. scutata,
being less robust posteriorly than in A.
calva. As in the other forms, it is narrower
anteriorly than posteriorly. The ventral
border is relatively straight, while that of
the other forms is posteriorly convex. Infra-
orbitals 2, 3, and 4 have not been identified.
The vomerine tooth patch in A. uintaen-
sis, as in A. fragosa, extends inore posteri-
orly than in A. calva (Fig. 19). The
\^omerine teeth are sharp and greatly curved
posteriorly; they exceed those of A. fragosa
and A. calva in number, each vomer bear-
ing between 40-50 teeth, as compared to
half that number in A. fragosa and A. calva.
The rostral and antorbital are identical to
that of the other species.
Brancliiocranium. The suture between
the anterior and posterior dermopalatine
cannot be discerned. In A. uintaensis the
dermopalatine has about twice the number
of teeth as in A. calva, and the tooth patch
extends more distad. The teeth are sharply
pointed, as are the vomerine teeth.
The hyomandibular is more deeply ex-
cavated between the opercular process and
the extensor ( dorsal ) surface, and the oper-
cular process is more massive and extends
further ventrad, forming a larger articula-
tion surface, as compared with the other
species of Amia. The articular surface of
the quadrate is more robust than in other
species of Amia and displays three cristae
ventrally rather tlian dorsally as in A. calva
and A. rolmsta (Janot, 1967: 144). The
ceratohyal resembles that of A. calva and
A. fragosa with the exception of its being
thicker at the neck of the proximal end.
The metapterygoid in A. uintaensis con-
forms very closely to that of A. calva in
outline and in the position of the anterior
Fig. 19. Comparison of vomers of A, Amia calva; B, A.
uintaensis; and C, A. fragosa.
52 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
basal process and the posterolateral otic
process.
The maxilla in A. uintaensis is more
robust and relatively longer, and its pos-
terior border is dorsoventrally wider than
in the other forms, particularly A. fragosa.
As in A. calva the small supramaxillary
notch occurs more anteriorly than in A.
fragoso. The dorsoposterior border is
rounded, as in A. calva and A. scutata,
rather than sharply angular, as in A. fragosa.
Anteriorly the maxilla is deeper and more
thickly ossified than in the other forms,
but this may be a function of greater size.
The supramaxilla resembles that of A. calva
in general morphology, being elongated and
narrowly tapered anteriorly, with a smooth-
ly rounded posterior end conforming to the
curve of the maxilla. The maxillo-supra-
maxillary suture is straight as in A. calva.
The premaxilla is identical to that of the
other species.
The dentary of A. uintaensis is similar to
that of A. calva and A. scutata in lacking
the dorsal shelf of the anterior lingual bor-
der of the alveolar ridge which occurs in
A. fragosa. The coronoids articulate more
or less vertically on the alveolar ridge, as in
A. scutata and A. calva. The anterodorsal
region of the dentary slightly overlaps the
ventral half, but not to the extent that it
does in A. fragosa; A. uintaensis seems to
be intermediate between A. fragosa and A.
calva in this feature, the latter having no
such ventral overlapping at the symphyseal
edge. The coronoid articulation surface of
the A. uintaensis dentary is thicker than in
A. fragosa and A. calva, but only slightly
more so than in A. scutata. At the termina-
tion of this surface, this thickened area of
bone forms the dorsal wall of the Meckelian
groove, as in A. calva. The ventral wall of
this groove is less well defined than in A.
calva, witli A. scutata being intermediate.
The anterior half of the dentary length in
A. uintaensis is evenly tapered to the
symphyseal edge; it is elongated and lacks
the sharp curve present in A. fragosa at the
midpoint of the alveolar ridge (Fig. 18).
There is only a trace of such a curve in the
dentaries of A. calva and A. scutata which
are also more elongated and evenly tapered
than in A. fragosa, although not to the ex-
tent that they are in A. uintaensis. Anteri-
orly, the bone is also relatively thicker than
in A. fragosa and A. calva; A. scutata also
displays this greater ossification at the
anterior end of the dentary. Posteriorly,
the dentary is very similar to that of A.
calva. The coronoid teeth are sharp and
conelike, extending to the midpoint of the
lingual surface, as in A. calva. As Janot
(1967) shows for A. robusta, the alveolar
ridge is more horizontal in A. uintaensis and
forms a more acute angle with the exterior
surface of the dentary than it does in A.
fragosa or A. calva; A. scutata is interme-
diate between A. uintaensis and A. calva in
this feature (Fig. 18). In A. uintaensis the
first coronoid (symphyseal) overlies only
the dorsal half of the anterior articular sur-
face of the dentary, as in A. calva and A.
scutata. The teeth are more sharply pointed
than in any of the other forms (Fig. 18).
The second coronoid is fragmentary, but
appears to resemble that of A. calva with
the exception of its having more sharply
pointed teeth. The prearticular specimens
available are fragmentary, but the lingual
surface possesses blunt-conical teeth similar
to those in A. calva and A. fragosa. Bor-
sally, however, these teeth are as sharply
pointed as the coronoid teeth. The angular
is slightly longer and higher than that of
A. calva. The posterior border is more ver-
tical, with the articular notch less pro-
nounced. It is more heavily ossified than
in A. calva, but this may be a function of
size. The surangular in A. uintaensis is
basically similar to that of A. calva, al-
though it is situated more dorsally and is
more rounded at the dorsal edge.
The gular is longer than that of A. calva
and A. fragosa (Fig. 20). It is also slightly
narrower at the posterior end than the
anterior end, while the reverse is generally
true in A. calva. Otherwise, the gular
strongly resembles that of A. calva. Despite
a few minor dissimilarities, the preopercu-
lum resembles that of A. calva. There is a
Fossil Amiids • Boreske 53
Fig. 20. Comparison of gulars of A, Amia calva; B,
A. uintaensis; and C, A. fragosa.
slightly more pronounced concavity in the
ventroposterior border than is exhibited in
A. calva; this concavity is altogether lacking
in A. fragosa. The line of curvature is about
the same as in A. calva; in A. fragosa the
preoperculum is more deeply curved. The
dorsal half is not quite as wide as the
ventral half, while in A. calva both ends are
fairly equal. In A. fragosa, however, the
dorsal half is much narrower and more
tapered than the ventral half, which is rela-
tively wider and bulbous. The operculum
in A. uintaensis is similar to that of A.
calva and A. scutata in operculum-depth/
operculum-length (Table 7). The suboper-
culum conforms in general morphology with
that of A. calva, although it is slightly more
robust, particularly in the posterior region.
The corners tend to be angular, as in A.
scutata and A. calva, rather than rounded,
as in A. fragosa. The interoperculum is
similar to that of A. calva, although more
robust. The anterodorsal border is more
convex than in A. calva, and is more deeply
impressed into the preoperculum. The
anteroventral border is narrowly tapered as
in A. calva, rather than smoothly rounded as
in A. fragosa. The first branchiostegal ray
conforms to that of the other species. Al-
though the lack of articulated material
makes any count of the rays difficult, in
MCZ 12916 there are 12 disarticulated
branchiostegal rays on the right side of the
cranial roof. As in A. fragosa the distal ends
of the rays are consistently rounded, rather
than squared as in A. calva.
Post-cranial skeleton. The supracleithrum
in A. uintaensis resembles that of A. calva
and A. fragosa, excepting the dorsal articu-
lation surface, which is rectilinear rather
than pointed as in A. calva. The distal
lateral border in the Paleocene specimens
lacks the notch that occurs in A. calva, but
this notch is present in the Eocene speci-
mens. The metacleithrum in A. uintaensis
is more elongated than in A. calva and A.
fragosa. The dorsal end is narrower than
in A. calva, and the ventral end is sciuared
off. The cleithrum in A. uintaensis is largely
similar to that of the other Ainia species,
but is more massive at the proximal end
than in A. calva, and the dermal sculpture
covers a greater area than in A. calva, ex-
tending to the distal border as in A. fragosa
and A. scutata (Fig. 21). The mid-distal
border is smoothly convex and lacks the
notch ventral to the metacleithrum which
is present in A. calva.
The preceding study of the vertebral
skeleton of A. calva revealed changes in
height/ width proportions, position of
chordal foramen, configuration of neural
and aortal facets, and in the basapophyscal
angles and length of basapophyses which
may be used here to discern similar trends
in A. uintaensis centra, for the fossil verte-
brae display the same features characteris-
tic of the Recent species even in disarticu-
lated state.
CM 25362 from the Bridger Formation is
the only specimen that has a relatively
complete, disarticulated, undistorted verte-
B
Fig. 21. Comparison of cleithra of A, Amia calva; B,
A. scufata; C, A. uintaensis; and D, A. fragosa.
54 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
bral column; as the centra are separable this
specimen is useful in comparisons with iso-
lated vertebrae. There are 59 centra pres-
ent: 25 trunk centra and 34 caudal centra,
including two fused urals. Many of the
preserved caudal centra are only fragments.
Since the articulated specimen (PU 13865)
has 85 vertebrae (see Table 9 for regional
numbers) it may be assumed that about 25
vertebrae are missing from CM 25362.
When comparing vertebrae from different
regions of the column in the two specimens,
it appears that CM 25362 lacks approxi-
mately six trunk and approximately twenty
caudal centra. The first anterior trunk cen-
trum present in the CM 25362 series pos-
sesses aortal facet configurations similar to
those of the seventh vertebra of the articu-
lated specimen ( PU 13865 ) . An articulated
but separable series of six uncrushed an-
terior trunk vertebrae (PU 10101), also
from the Bridger Formation, aids in the
reconstruction of the anterior region of the
A. uintaensis vertebral column (Figs. 22-
25). The basapophyseal angles of these six
PU 10101 vertebrae do not vary from 180
degrees. The first six anterior trunk verte-
brae from a partly disarticulated vertebral
column from the Paleocene specimen (PU
16236) resemble the six PU 10101 centra in
length and shape of aortal facets, even
though PU 16236 is a smaller individual.
The height/ width ratio of these latter centra
is difficult to determine, however, since the
specimen underwent postdepositional crush-
ing. The nearly complete vertebral series
of the CM 25362 specimen has been used
for the construction of the remaining trunk
and caudal region in the model of the A.
uintaensis vertebral column. The trunk
centra of CM 25362 have been arranged
according to basapophyseal angles that de-
crease from 180 to 46 degrees, as in A. calva.
Decreasing size was used to arrange the
caudal vertebrae.
Although A. uintaensis occurs much ear-
Table 14. Angle of basapophyses, length, height, and width of vertebrae of Amia
uintaensis compared with type specimens of synonymized taxa as illustrated
IN figure 22
Relative
Vertebral
Number
Specimen
Angle of
Basapophyses
( Degrees )
Length
(mm)
Height
(mm)
Width
(mm)
6
A. uintaensis PU 10101
A. uintaensis PU 16236
P. uintaensis ANSP 8044
P.'sp. USNM 170973
A. uintaensis PU 10101
A. uintaensis PU 16236
A. whiteavesiarm NMC 6197
P. sp. FMNH P27465
A. uintaensis PU 10101
A. uintaensis PU 16236
P. sp. USNM 3966
P. medius USNM 3959
A. uintaensis PU 10101
A. uintaensis PU 16236
A. uintaensis PU 10101
A. uintaensis PU 16236
A. uintaensis PU 13865
A. uintaensis PU 10101
A. uintaensis PU 16236
P. uintaensis ANSP 5558
ISO"
180°
180°
180°
180°
8.0
32.0
45.0
6.0
31.0
44.0
8.0
32.0
46.0
5.5
19.0
29.0
8.0
32.0
44.5
8.5
28.5
39.0
8.5
29.0
40.0
9.0
28.0
36.0
10.0
33.0
43.0
8.0
30.0
41.5
8.5
21.5
29.0
8.5
22.0
30.0
10.0
33.0
44.0
7.5
33.0
40.8
11.0
33.5
44.0
9.5
31.5
39.0
4.5
16.5
21.5
11.0
34.0
42.5
9.0
33.5
34.0
10.5
32.5
40.0
Fossil Amiids • Boreske 55
licr in time than A. calva and A. scutata, it liodicd form tlian its contemporary, A.
has approximately the same total number of fra^osa, which has a mean of 65 centra,
centra (85), and like them is a longer- Tlu> vertebral column of A. umfaenm does,
B
4. uintaensis
PU lOIOI
A. uintaensis
PU 16236
P. uintaensis
ANSP 8044
A . whiiteavesiana
NMC 6197
R sp.
USNM 3966
A. uintaensis
PU 13865
P. sp.
USNM 170973
P sp.
FMNH P27465
R uintaensis
ANSP 5558
P medius
USNM 3959
Fig. 22. First anterior trunk vertebrae (A,B) of Amia uinfaens'is compared with type specimens of synonymized taxa
(refer to Table 14 for data).
56
Bulletin Museum of Comparative Zoologij, Vol. 146, No. 1
11
R sp. P. sp.
USNM 170973 USNM 3962
P. Sp.
USNM 170973
"-^,/
P. plicotus
AMNH2539
P sp.
PU 20523
P Sp.
USNM 170973
P sp.
USNM 170973
12C
12d
i
Amia. sp.
ANSP2337
P. plicafus
USNM 170974
P. medius
USNM 3959
12a
p. medius P. sp.
YPM 6238 FMNH P27465
P. medius
USNM 3959
12b
Amia sp.
ANSP 2339
Fig. 23. Seventh through fourteenth mid-trunk vertebrae of Amia uinfaensis compared with type specimens of
synonymized taxa (refer to Table 15 for data).
Fossil Amiids • Borcske 57
however, differ meristieally from that of 36 (mean) in A. sctitata. The number of
A. calva and A. scutata in number of verte- diplospondylous vertebrae is 20-21, as eom-
brae in the various regions. There are 31 pared with 14-17 in A. calva and 15 in
trunk eentra in A. uintacnsis (PU 13865), A. scutata. This variation from A. ra/i;« and
as opposed to 37 (mean) in A. calva and A. scutata in the organization of the verte-
15
/? plicotus P. medius
USNM 3958 USNM 3959
R medius
YPM 6239
R sp.
USNM 3966
P symphysis
PU 10099
19a
19b
P medius
USNM 3959
R sp.
USNM 3966
R sp
USNM 3962
P sp. P sp.
USNM 3966 FMNH P27465
P medius
YPM 6240
P sp.
USNM 3966
19C
R sp. R sp-
FMNH P27465 USNM 3963
P. sp. P medius
USNM 3966 USNM 3959
R medius
USNM 3959
R loevis
USNM 3968
P uintaensis
ANSP 3151
Fig. 24. Fifteenth through twenty-second posterior trunk vertebrae of Amio uiniaemis compared with type speci-
mens of synonymized taxa (refer to Table 16 for data).
5(S Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
26
27
28
29
30
31
Psp.
YPM 6242
/? uintaensis
ANSP3I5I
P. loevis
PU 10109
R laevis
PU 10109
P. media
ANSP5632
P Sp.
USNM 3963
Amia sp.
ANSP2338
P symphysis
PUIOIIO
A. mocrospondyla
NMC 6198
34
37
39
43
44
45
50
51
59
m
P. laevis
USNM 3968
P sp.
FMNH PF4509
P sp.
USNM 3966
P. medius
USNM 3959
P. medius
USNM 3959
P sp.
USNM 5450
P. medius
USNM 3959
P. corsoni
USNM 3961
Fig. 25. Posterior trunk and caudal vertebrae of Am/o umfoensis compared with type specimens of synonymized
taxa (refer to Table 17 for data).
Fossil Amiids • Boreske 59
Table 15. Angle of hasapophyses, length, height, and width ok vehtehuae of Amia
iiintacn.sis compared with type specimens of synonymized taxa as illustrated
IN figuhe 23
Relative
Vertebral
Number
Specimen
Angle of
Basapophyscs
( Def^rees )
Length
(mm )
Height
(mm)
Width
( mm )
7
8
9
11
12a
12b
12c
12d
14
P. sp. USNM 170973
P. sp. USNM 3962
P. sp. USNM 170973
P. plicatus AMNH 2539
P. sp. USNM 170973
P. .s7>. PU 20523
P. sp. USNM 170973
P. 77ic(lius YPM 6238
P. sp. FMNH P27465
P. mcdius USNM 3959
Aiiiiu sp. ANSP 2339
A7nia sp. ANSP 2337
P. ])licatus USNM 170974
P. mcdius USNM 3959
179°
8.5
19.5
25.5
7.0
20.0
28.0
178°
8.5
19.5"
26.0
177°
6.0"
18.0^
24.0
8.0
21.0
25.5
174°
7.5
25.0
30.0
7.5
22.0'
25.0
6.0
19.0
24.0
171°
11.0
30.0
35.0
9.0
22.0
28.0
167°
10.0
29.0"
40.0
166°
10.0
29.0
38.5
163°
8.0
24.0
30.0
160°
7.5
19.0
23.0
Est.
Taulk 16. Angle of hasapophyses, ijiingth, hkic;ht, and width of verteijuae of Amu/
uintactisis compared with type specimens of synonymized taxa as illustrated
IN figure 24
Relative
Vertebral
Number
15
17
18
19
19a
19b
19c
20
21
22
Sjiccimen
P. plicatus USNM 3958
P. mcdius IfSNM 3959
P. mcdius YPM 6239
P. sp. USNM 3966
P. sijmphysis PU 10099
P. mcdius USNM 3959
P. sp. USNM 3962
P. sp. USNM 3966
P. ,v/;. USNM 3966
P. sp. FMNII P27465
P. i7icdius YPM 6240
P. sp. USNM 3966
P. sp. FMNH P27465
P. sp. USNM 3963
P. sp. USNM 3966
P. Tticdius USNM 3959
P. mtY/tt/.v USNM 3959
P. /at't;i.v USNM 3968
P. uiiitacrisis ANSP 3151
Anglo of
Basapophyses
( Degrees )
Length
(mm)
Height
(mm)
Width
( mm )
156°
8.5
8.5
8.5
8.5
5.5
22.5
23.0
22.0
21.0
15.5
25.0
27.0
24.5
25.5
20.0
153°
9.0
9.0
24.0
23.0
29.0
28.0
149°
9.0
21.0
25.5
143°
8.5
11.0
23.0
28.0
27.8
33.0
139°
7.0
19.0
23.0
138°
—
—
136°
11.0
7.0
25.0
16.5
35.0
20.0
132°
9.0
9.0
21.0
22.0
27.0
26.0
122°
9.5
22.0
27.0
117°
10.0
12.0
26.0
28.0
30.0
29.0
60
Bulletin Museum of Coniparative Zoology, Vol. 146, No. 1
Table 17. Angle of basapophyses, length, height, and width of
uintaensis compared with type specimens of synonymized taxa
IN figure 25
vertebrae of Amia
as illustrated
Relative
Vertebral
Number
Specimen
Angle of
Basapophyses
( Degrees )
Length
( mm )
Height
( mm )
Width
( mm )
24
P. sp. YPM 6242
102°
9.0
17.0
22.0
26
P. tiintaensis ANSP 3151
97°
10.0
23.0
28.0
27
P. laevis PU 10109
90°
13.0
29.0
33.0
28
P. laevis PU 10109
83°
14.0
28.0
30.0
29
P. media ANSP 5632
80°
8.0
16.0
18.0
30
P. sp. USNM 3963
P. symphijsis PU 10110
62°
11.0
6.0
14.0
13.5
31
Amia sp. ANSP 2338
A. macrospondyla NMC 6198
46°
13.0
12.0
26.0
25.0
23.0
22.0
34
P. laevis USNM 3968
6.5
22.0
18.0
39
P. sp. FMNH PF 4509
7.0
19.0
18.0
43
P. s/;. USNM 3966
7.0
19.0
18.0
44
P. medins USNM 3959
6.0
16.0
18.0
45
P. 7ne(/ii« USNM 3959
5.5
17.5
18.0
50
P. s/;. USNM 5450
7.0
15.0
13.5
51
P. »!«//»« USNM 3959
5.0
17.0
11.0
59
P. corsonii USNM 3961
4.0
11.0
10.0
bral column into region.s and types of verte-
brae appears to be a useful taxonomic
character of A. nintaensi.^.
The neural, aortal, and haemal facets do
not appear to vary much from those of
A. calva. The first six ventral aortal facets
show basically the same pattern for both
species (Figs. 11, 22). The angle of basa-
pophyses in A. uintaensis differs from that
of A. calva in two ways. The first six verte-
brae all have basapophyseal angles of 180
degrees, and it is not until the seventh
vertebra that these angles gradually begin
to decrease. Because of this more posterior
beginning in the decrease of the angles and
because there are fewer tinmk vertebrae, the
rate of decrease of the basapophyseal angle
is greater. These angles range from ISO
degrees anteriorly to approximately 45 de-
grees posteriorly, about the same as the
range for A. calva.
The intracolumnar variation in centrum
shape seen in the vertebral column of Re-
cent A. calva also occurs in A. uintaensis
( Fig. 14). In some respects the latter shares
certain characteristics with A. calva. The
first centrum is broad and thin, and usually
lacks basapophyses (Fig. 22). However,
centra between the fourth and twentieth
vertebrae begin to acquire an almost sub-
triangular outline, as opposed to the sub-
elliptical form of the A. calva trunk centra
(Fig. 12). The subtriangular shape may be
a function of greater size of the centra.
The chordal foramen is open in all known
vertebrae of A. uintaensis from the Paleo-
cene. Eocene, and Oligocene, but is often
filled with detritus during fossilization.
Estes (1964: 42) observed that Cretaceous
specimens as well as the Late Faleocene
specimen (PU 16236) had the chordal
foramen smoothly closed with bone. A re-
Fossil Amiids • Boreske
61
examination of PU 16236 reveals that the
chordal foramen is actually filled with fine
sediment rather than bone, so that the
character of the closed foramen can only be
applied to the Cretaceous specimens.
Chordal foramen position in all specimens
shows slight intracolumnar variation along
the tiimk as in A. calva, although occurring
more dorsally. In the caudal region there is
virtually no difference between the two
forms.
Leidy characterized "Protamia" uintaensis
on the basis of five centra and one basioc-
cipital. His height/ width proportions were
described in relation to those in an un-
diagnostic intracolumnar standardization of
the centra of the A. calva vertebral column.
My measurements of the anterior trunk
centra reveal that the holotypc ANSP 5558
has a width 1.3 times the height, and para-
type ANSP 8044 has a width 1.6 times the
height. Other paratype centra are posterior
trunk centi-a with width/ height ratios of
approximately 1:1. Romer and Fryxell
(1928: 521) described a displaced posterior
trunk centrum as having a height of 10 mm
and a width of 12.5 mm, about the same as
in ANSP 5558. Estes (1964: 43), in his
discussion of the height/ width proportions
of A. uintaensis centra, misinterpreted
Leidy 's (1873a, 1873b) diagnosis of "Proto-
mia' uintaensis and Romer and Fryxell's
(1928) diagnosis of "Paramiatus ^wleyij'
indicating that vertebrae of tlie former were
three times as wide as deep, those of the
latter two times. Estes was correct, how-
ever, in his assumption that there is intra-
columnar variation in height/ width ratios.
The general pattern of intracolumnar
variation in the A. uintaensis vertebral
column is quite similar to that of A. calva;
there is the same trend from horizontally
elliptical centra to circular or vertically
elliptical centra ( Fig. 14 ) . Thus the earlier
diagnoses of A. uintaensis using height/
width ratios that attributed the proportions
of the anteriormost trunk vertebrae to the
entire column are undiagnostic.
On the basis of isolated centra and skull
material, the most commonly used character
in differentiating A. uintaensis from A. calva
has been the former's greater size. How-
ever, the articulated specimen (PU 13865),
which is the smallest known A. uintaensis,
is only 146 mm longer than A. fra^osa
(FMNH 2201) and 16 mm longer than the
largest A. calva known to me (UMMZ
197683). Estes (1964) suggested that the
widening of the A. uintaensis vertebrae
might be a function of its greater size;
Gould's (1966) statement that internal
elements generally increase at allometric
rates to provide sufficient surface area to
maintain the external surface area offers a
partial explanation as to why the large A.
uintaensis vertebrae have greater width in
proportion to height than they do in smaller
amiid vertebrae.
Discussion
Two species of Protamia, one species of
Ilypamia, six species of Pappichthys, and
three species of Amia have been described
on vertebral characters from isolated centra
and disarticulated cranial elements (Table
19). With the exception of Amia ichiteaves-
iana, A. selwyniana, and A. macrosponclyla
from the Oligoeene Cypress Hills Forma-
tion of Alberta, all these taxa are based on
material from the Bridger Basin, Bridger
Formation, of Wyoming. Each of these 12
taxa will be re-evaluated in the following
discussion. Of the twelve species and four
genera, 'Trotamia" uintaensis (Leidy,
1873a) is the oldest name. Leidy's type
specimens are all trunk vertebrae. The
holotypc ANSP 5558 (Fig. 22) is approxi-
mately the sixth anterior vertebra and
displays the characteristic subtriangular
outline of other specimens. The paratypes
include trunk vertebrae ( ANSP 8044, 3151 ),
and a large basioccipital (ANSP 5622). The
holotypc vertebrae and the basioccipital
are considered diagnostic for Amia uintaen-
sis, on the basis of their possessing the
characteristic subtriangular vertebral out-
line, and a kidney-shaped articular surface
of the basioccipital.
Leidy (1873a) described Protamia media
from two large trunk centra from the
62 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
Bridger Formation of Wyoming. His main
criterion for distinguishing this form from
A. calva and from the other species of
"Protarnia" was that the vertebrae were
twice the size of A. calva vertebrae and
"somewhat smaller than Protamia uintaen-
sis" (Leidy, 1873b). The holotype USNM
2181 appears to be from the anterior trunk
region ( approximately the seventh or eighth
centrum, as suggested by its proportions
and configurations of aortal facets). The
basapophyseal angle is approximately 178-
180 degrees. The paratype ANSP 5632 is
from the posterior trunk region, with an 80-
degree basapophyseal angle, which is ap-
proximately equivalent to the twenty-ninth
centrum in A. uintaensis (Table 17; Fig.
25). Cope ( 1884, plate 4, figs. 7-20) figured
"PappiclitJiys mcdius" on the basis of 14
disarticulated centra from the same locality
(USNM 3959). Eight of these are from the
trunk region and correspond to centra
within the anterior to mid-trunk region of
A. uintaensis (Tables 15-16; Figs. 23-24).
The remaining six centra correspond to cen-
tra in the caudal region (Tables 16-17;
Figs. 24-25). Cope gave no description, but
in figuring these specimens he allocated to
them his own genus, emending Leidy's
(1873a) prior nomenclature. Both Leidy
and Cope had apparently assumed that the
characteristics of one or a few vertebrae
represented those of the entire coliunn.
Both species fall well within the size range
of A. uintaensis (Tables 14-17), and are
here considered synonyms of the latter.
Leidy (1873a) describc-d Uypamia ele-
gans from one small trunk vertebra. He
characterized this form as possessing a cen-
trum that was characteristically "short in
proportion with its breadth, and it presents
sutural impressions for a contiguous pair of
neural arches" (Leidy, 1873b). ANSP 5580
appears to be from the mid-trunk region,
comparable to approximately the nineteenth
centrum as suggested by its proportions and
configuration of aortal facets. The basapo-
physeal angle is 138-139 degrees. These
character-states and the small size are not
unique, occurring as the do in all the other
species of Amia; Hypamia elegans is there-
fore a nomen duhium.
Cope (1873) described Pappiclithys
plicatus from the anterior portion of a large
left dentary (AMNH 2539). Other type
material included two premaxillae, a right
quadrate, a left epihyal, an anterior portion
of an ectopterygoid, three trunk vertebrae,
and numerous fragments of angulars. He
characterized this form primarily on the
basis of dermal sculpture of the "cranial
fragments being roughly grooved." The
angular in A. uintaensis is generally
marked by more pronounced dermal sculp-
tiu-e than the other mandible elements.
His diagnosis of the vertebrae (USNM
3958) is based on proportions and mor-
phology of neural and aortal facets, both
of which correspond to various trunk ver-
tebrae in A. uintaensis (Tables 15-16;
Figs. 23-24). The description of the re-
maining elements conforms with other
elements of A. uintaensis. Pappichthys
plicatus is tlierefore a synonym of the latter.
Cope (1873) described Pappichthys
sclerops from a large left dentary. He
characterized this form as possessing a
dentary "more compressed and deeper" than
that in A. calva and other species of "Pap-
pichthys." The dentary (USNM 3965) in
all respects greatly resembles all dentaries
that have been referred to A. uintaensis, and
I regard Pappichthys sclerops as a synonym
of the latter.
('ope (1873) described Pappichthys
laevis from a large anterior dentary frag-
ment (USNM 3968). Other type materials
include a premaxillary fragment, fragments
of angulars (AMNH 2570), a left quadrate
fragment, a trunk vertebra fragment, and a
caudal vertebra. Although Cope distin-
guished this taxon from other species of
Pappichthys on vertebral proportions, vari-
ances in dermal sculpture, dentary alveolar
count, and obliqueness of alveolar face,
these character-states occur in A. uintaensis.
PappicJitJujs laevis is therefore a synonym
of the latter.
»
Fossil Amiids • Boreske
63
Cope (1873) described Pappichthys sym-
pJu/sis from two large fragments of trunk-
vertebrae and a iiral (USNM 3960). His
diagnosis rests primarily on eonfiguration
of neural faeets and basapophyseal length.
Osborn et cil ( 1878: 104) later reported two
eaudal vertebrae as cotypes (PU 10099,
10110). Cope (1873) described Pappich-
thys corsonii from 12 centra (USNM 5475-
5476), a basioccipital (USNM 5476), and
a left dentary fragment (USNM 3961). He
distinguished this form from Pappiclithys
sympliysis on different neural facet mor-
phology, basapophyseal length, and height/
width proportions. Merrill (1907: 14) cites
^'PappicJitJujs sympliysis = Pappichthys cor-
sonii' without further discussion. The cen-
tra of both forms conform to centra in the
vertebral column of A. uintaensis (Table
17; Fig. 25) and the characters assigned to
the dentary and basioccipital of Pappich-
thys corsonii are also found in A. uintaensis;
thus both P. sympliysis and P. corsonii arc
synonyms of A. uintaensis.
From the Early Oligocene Cypress Hills
Formation, Saskatchewan, Cope (1891)
described Amia wliiteavesiana from an an-
terior vertebra (NMC 6197), and Amia
macrospondyJa from a caudal vertebra
(NMC 6198). Both these forms were
founded on variations of vertebral charac-
ters (height/ width proportions, lack of
basapophyses, and chordal foramen posi-
tion) that are also represented in the verte-
bral column of A. uintaensis. The type
centrum of A. tiJiiteavesiana corresponds
approximately to the second anterior verte-
bra in A. uintaensis (Table 14; Fig. 22),
that of the type centrum of A. macrospon-
(Jyla with the thirty-first centrimi in A.
uintaensis (Table 17; Fig. 25). Prior to
the appearance of Cope's ( 1891 ) publica-
tion. Ami (1891), in his review of the
Cypress Hill fauna, mistakenly listed A.
whiteavesiana under the name A. selwyni-
ana. A. macrospondyhi and A. ivhiteavesi-
ana are here considered synonyms of A.
uintaensis; A. sehcyniana is a iiornen
nudum.
Comments on European and
Asian Forms
Janot ( 1967 ) described a large amiid,
Amia rohusta, from the Late Paleocene of
France, on the basis of disarticulated
material. She distinguished this form from
A. calva and A. russeUi on the angle of the
ventral border of the dentary face, and on
morphology of the parasphenoid tooth-
bearing surface in addition to other minor
morphological differences. Many of the di-
agnostic elements or associations on which
A. uintaensis is based, such as coronoid and
vomerine teeth, regional vertebral counts
and dorsal cranial elements, are missing in
her material. The elements that she does
figiue, however, closely resemble the com-
parative bones in A. uintaensis. Simi-
larities include rounded distal ends of
branchiostegal rays (also in A. fragosa),
subtriangular morphology of trunk verte-
brae, extensive surface of parasphenoid
tooth-patch, and shallow orbital notch in
frontal (also in A. scutata and A. calva).
These marked similarities suggest that A.
rohusta is a synonym of A. uintaensis.
Current work on the relationship of the
North American and European continents
in the Early Cenozoic ( McKenna, 1972)
indicates that they were connected until tlie
Early Eocene and that there is great sim-
ilarity between the Paleocene and Early
Eocene mammalian taxa at that time.
There is thus no zoogeographic problem in-
herent in synonymizing these two species.
Hussakof (1932) described Pappichthys
mongoliensis from disarticulated elements
from the Late Eocene Ulan Shireh beds of
the Shara Murun region. Inner Mongolia
(collected by the American Museum Cen-
tral Asiatic Expeditions.) At the time of
Hussakof's description, this collection
(AMNH 6372) represented the most exten-
sive material of ''Pappichthys." The collec-
tion includes numerous dentaries, maxillae,
three gulars, three opercula, three cleithra,
an hyomandibular, a supracleithrum, a
vomer, and trunk and caudal vertebrae.
64 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
Hussakof distinguished this form from A.
calva by the length of the dentaries and
the morphology of the operculum, and
from species of "Pappichthys" and "Pro-
tamia" on the basis of comparison of verte-
bral size. A comparison of the Mongolian
material with A. uintaensis shows some dis-
similarities, but there is still a closer affinity
between this form and A. uintaensis than
with the other species of Amia. The vomer
bears numerous sharp vomerine teeth; the
hyomandibular is deeply arched, and the
lingual face of the dentaries conforms to
that of A. uintaensis. The dentary, however,
is quite elongated anteriorly, the supra-
cleithrum is narrower, and the dorsal
border of the operculum is short and
ascends at a 30-degree angle rather than
being horizontal as in A. uintaensis (and
in other Amia species). The extrascapular
is narrow and tapered to a point rather than
flattened medially. Thus, although Pappich-
thys mongoliensis is similar to A. uintaensis
in many features and is clearly related to it,
it also differs in some respects. It un-
doubtedly belongs to the genus Amia, and
retention of all the Mongolian specimens in
Amia mongoliensis seems the most practical
alternative at this time. The Mongolian
higher vertebrate taxa indicate that the
Turgai Straits at least partially isolated
Mongolia from Europe during at least part
of the Cretaceous, Paleocene, and Eocene,
and that probably little exchange took place
until the Late Eocene (Szalay and Mc-
Kenna, 1971: 280-281). It may be possible
that A. mongoliensis evolved from A.
uintaensis during this migration.
Amia cf. uintaensis
Hypodigm. Cretaceous. Lance Forma-
tion, Wyoming: CM 256, YPM 6311, trunk
vertebrae; UCMP 56276, two fragments of
a single vertebra; UCMP 56277, one com-
plete vertebra, one vertebral fragment, one
left maxillary fragment. Hell Creek Forma-
tion, Montana: AMNH 6385, trunk vertebra;
MCZ 9334, dentary tooth tips. Aguja For-
mation, Texas: UMM collections, maxillary
fragment. Ojo Alamo Formation, New
Mexico: USNM collections, trunk vertebra.
Discussion
Cretaceous specimens of large amiids
occur in both Lance and Hell Creek for-
mations and consist mostly of isolated
and broken centra, and teeth that have
been identified primarily on the basis of
size. The characteristic subtriangular out-
line of the trunk vertebrae is even more pro-
nounced in these Cretaceous specimens,
wherein the lateral centrum walls between
the basapophyses and the aortal facets are
concave ( Fig. 26 ) . The chordal foramen is,
as Estes ( 1964: 42) noted, closed with bone,
as are one-third of the vertebrae referred to
A. fragosa from the Lance Formation. How-
ever, Estes observed lateral concavities be-
tween the neural facets and basapophyses
in a large vertebral centrum (AMNH
6385) from the Hell Creek Formation
(mistakenly cited by him as AMNH 6835
from the Oldman Formation of Alberta).
Estes apparently confused neural with
aortal facets and thus figured the vertebra
upside down. Correct orientation of the
centrum (Fig. 26) shows concavities be-
tween the basapophyses and the aortal
facets. Thus, Estes was incorrect in con-
cluding that A. fragosa, A. calva, and the
Eocene specimens of A. uintaensis "also
seem to lack the concavity between the
'basapophysis' and neural arch present in
the large Cretaceous specimens." Two
other specimens from the Lance Formation
(YPM 6311, CM 256; Fig. 26) also show
the prominent concavities between the
basapophyses and aortal, rather than neu-
ral, facets. In addition to the vertebrae,
Estes described a maxillary fragment as
being larger and more robust than that of
A. fragosa, although "characteristically
amiid in tooth implantation and general
shape." A more complete maxillary frag-
ment (UMM collections) from the Aguja
Formation ( Big Bend National Park,
Brewster County, Texas) conforms with
Estes' (1964) description.
Fossil Amiids • Borcnke 65
B
Hlii
H
Fig. 26. Comparison of different Cretaceous vertebrae. Am'ia cf. uinfaensh: A, anterior trunk vertebra, CM 256,
Lance Formation, Wyoming; B, posterior trunk vertebra, AMNH 6385, Hell Creek Formation, Montana; C, mid-trunk
vertebra, YPM 6311, Lance Formation, Wyoming. Chondrichthyes: D, E, G (thin section), trunk vertebrae, FHKSCM
13024-9, Black Creek Formation, North Carolina; F, trunk vertebra, MCZ 12879, Peedee Formation, North Carolina.
Cetacean: H, caudal vertebra, FHKSCM 13025, Calvert Formation?, North Carolina.
1 rz dorsal, 2 ^ articular surface, 3 ::= ventral
66
Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
Only three new centra and a maxillary
fragment have been identified since Estes'
(1964) study. The vertebrae, as noted
above, differ in certain minor respects from
the Paleocene and Eocene specimens.
Whether or not this material actually repre-
sents A. uintaensis or an earlier stage of
evolution can only be determined when
more complete Cretaceous material is avail-
able.
Amia scutata Cope, 1875
Amia dictycephala Cope, 1875: 3.
Amia exilis Lambe, 1908: 12.
Holotijpe. USNM 5374, incomplete spec-
imen lacking the head and body anterior to
the middle of the dorsal fin; anal and part
of dorsal and caudal fins well preserved.
Type locality arul horizon. Florissant,
Colorado. East half of section 2, T 13 S,
R 71 W, Teller County, Colorado; Flor-
risant Formation.
Age Range. Chadronian (Early Oligo-
cene) to Orellan (Middle Oligocene).
Hypodigm. Oligocene. Cypress Hills
Formation, Saskatchewan: NMC 6200, 6205,
vertebrae; NMC 6201, basioccipital. Chad-
ron Formation, South Dakota: PU 17172,
left dentary with posterior coronoid bearing
teeth, and a trunk vertebra. Lower Brule
Formation, South Dakota and Nebraska:
FMNH PF4508, PF4509, CM 3814, verte-
brae; FMNH PF4506, right vomer bearing
teeth. Florissant Formation, Colorado: PU
10172, nearly complete specimen (counter-
part = YPM 6243, anterior half; USNM
4087, caudal half); YPM 6241, complete
caudal region (with counterpart); UMMP
V-57431, nearly complete specimen; USNM
3992, partial specimen, lacking skull and
tail; AMNH 2802, nearly complete skiill;
AMNH 2670, partial specimen, lacking
skull and caudal region; AMNH 2671,
caudal region.
Known distribution. South Dakota, Ne-
braska, Colorado, and Saskatchewan.
Revised diagnosis. Vertebral meristics
similar to those of A. calva, but head/
standard-length proportion is intermediate
between that of A. uintaensis and A. calva.
Extrascapular thicker at distal end than in
A. calva, with concave posterior border.
Pterotic more similar to that of A. uintaen-
sis than of A. calva; anterior portion narrow
and extended laterally to the frontal. Or-
bital excavations more marked than in A.
calva, but not as deep as in A. uintaensis or
A. fragosa. Preoperculum resembles that of
A. uintaensis more than that of A. calva,
being narrower dorsally than ventrally.
Symphyseal incurving of the dentary less
than in A. calva, but greater than in A.
uintaensis. Ventroposterior process of
cleithrum heavily sculptured as in A. fragosa
and A. uintaensis. Infraorbital 4 larger
than infraorbital 5 as in A. fragosa and A.
uintaensis. Ossification of cranial bones ex-
tensive as in other fossil species, greater
than in A. calva. Greatest known standard-
length 390 mm.
Introduction
Cope's (1875: 3) description of Amia
scutata is based on a specimen lacking the
head and body anterior to the middle of
the dorsal fin, from the Middle Oligocene
Florissant Fomiation near Florissant,
Colorado. He distinguished this form
from Amia dictyocephala (found in the
same deposit; Cope, 1875) and Amia calva
by its larger scales "of which only seven
and a half longitudinal rows are visible
above the vertebral column." Cope de-
scribed A. dictyocephala from two partially
complete specimens lacking skidls and
caudal fins (USNM 3992, AMNH 2670),
two complete caudal regions (AMNH 2671,
USNM 4087), and a nearly complete skull
(AMNH 2802); Osborn et al. (1878) later
described another specimen of A. scutata
from the same deposit. Tliis specimen
was more complete, consisting of an axial
skeleton and a crushed skull. They believed
A. scutata to be a valid form, differing from
A. calva in having a proportionately larger
head.
Comparison of known specimens of A.
scutata revealed that the counterparts to
the specimen described by Osborn et al.
( PU 10172 ) were separated and sold to two
Fossil Amiids • Borcskr 67
Fig. 27. A, Amia scutata UMMP V-57431; B, A. scufafa PU 10172; C, A. "dictyocephala" USNM 3392; D, A.
"dicfyocepbala" AMNH 2670.
different miiseiims. The caudal portion of and is one of the paratypes used by Cope
the counterpart was found in the National (1875) in his description of A. dictijo-
Museum of Natural History (USNM 4087) cephala. The anterior region was found
68
Bulletin Museum of Comparative Zoologij, Vol. 146, No. 1
unlabeled at the Yale Peabody Museum
(YPM 6243; Plate 4).i
In 1967 another nearly eompk^te speci-
men was discovered from the same deposit
(Fig. 27A) and Cavender (1970: 42) re-
ported the specimen A. dictyocepliala as
differing from A. calva in having a larger
infraorbital 4, in the sculptin-e of cleithrum,
and "by its proportionately larger head and
orbit, and somewhat shorter body."
Fossil Record
Other than the Florissant Formation, the
only deposits from which elements of A.
scutata can be identified are the Cypress
Hills Formation of Saskatchewan, Chadron
Formation of South Dakota, and the Lower
Brule Formation of South Dakota and
Nebraska. Becker (1961: 38) reported
amiid scales (UMMP collections) from
the Late Oligocene Passamari Formation
and Middle Oligocene Grant Horse Prairie
Shale of Montana (Becker, 1962). Since
no specific characters for scales of Amia
have yet been determined, it is best to
allocate this material to Amia sp. Skinner
et al. (1968: 415) has reported Amia .sp.
vertebrae (F:AM 42947) from the Early
Miocene Turtle Butte Formation of South
Dakota. Only two specimens were found;
since the vertebrae of A. scutata and A.
calva are morphologically and meristieally
similar. Skinner et al.'s identification is the
only possible one at this time. The strati-
graphic range of A. scutata is therefore lim-
ited to the Early and Middle Oligocene.
Description
Neurocranium.. The basioccipital (PU
10172, NMC 6201 ) is similar to that of A.
calva. The only available parasphenoid
(PU 10172) is poorly preserved, but closely
resembles that of A. calva in length and
position of ascending processes.
The extrascapular in A. scutata differs
slightly from that of A. calva in that the
distal end is relatively thicker and the
1 The counterparts ( USNM 4087, YPM 6243) to
PU 10172 have been .subse(iuently accjuired by the
Museum of Natural History, Princeton University.
L< orbital length
D- orbital depth
^a^dermosphenotlc angle
A. fragosa D/L=O.I76mn.
A. uintaensis D/L=O.I55mn.
18°
Z^"! 40 = I34<'
A scutata D/L=O.I32mn.
I5«
A calva D/L"O.IOOmn.
15°
IB"
40=145°
Z4»r 4.0=135°
A. cf. scutata O/L'0.121
[■0=137°
Fig. 28. Orbital dimensions of ^m\a spp.
posterior lappet is less pronounced; also,
the posterior border is more convex (Fig.
15). As in A. calva, however, the proximal
anterior corner is squared off, and the
medial suture is relatively long. The
pterotic in Ax. scutata resembles that of A.
uintaensis more than that of A. calva in
general morphology, since the anterior half
is narrower than the posterior half; in the
Recent species the ends are nearly sym-
metrical. The anterior border extends fur-
ther laterally than in A. calva, and, as in
A. uintaensis, adjoins the distal lateral side
of the frontal, rather than the posterior
border as in A. calva. The dermosphenotic,
parietal, frontal, and nasal of A. .scutata
conform to these bones in A. calva. The
parietal/ frontal ratio is marginally within
the lower limit of the range of A. calva
(Table 7). The orbital excavation in the
Imxssil Amuus • liorcske
69
frontal (Fit:;. 2cS) is greater than in A. caha
bnt less than in A. jruii^osa or A. uintaensis.
Snprascapulars, antorbitals, and rostrals are
not preserved.
The laerinial is similar to that of A. calva,
b(>arint!; a posterior noteh for the reeeption
of infraorbital 2, but in A. scutata the laeri-
nial is more robust. Infraorbital 2 and infra-
orbital 3 are similar to these bones in A.
calva. Infraorbital 4 is more massive pos-
teriorly than in A. calva; it exeeeds infra-
orbital 5 in dorsoventral length, and the
posterodorsal corner, which in A. calva is
markedly acute is, in A. .scutata, more
squared off. This bone more closely re-
sembles that of A. jraii^osa: it is not avail-
ablc> for comparison in A. uintacnsis. Infra-
orbital 5 is less massive posteriorly than in
A. calva; in this feature it resembles that of
A. fra'^osa. It is also, as in A. fra^osa and A.
uintacnsis, dec>per anteriorly than in A.
calva.
Branchiocranium. The supramaxilla in
A. scutata is elongated and tapered to a
point anteriorly, with a relatively straight
ventral border as in A. calva. It is slightly
longer and more robust posteriorly, the
posterodorsal border being higher and less
obliquely curved than in A. calva, A. uin-
tacnsis, and A. jra<^osa. The preniaxilla
resembles that of A. calva. The maxilla is
wider posteriorly and more ossified anteri-
orly than that of A. calva, but otherwise
agrees with the bone in the Recent .species.
Dermopalatine, autopalatine, entoptery-
goid, ectopterygoid, metapterygoid, and
vomer are not pr(\served. Ho\v(^ver, conical
vomerine teeth are displayed on PU 10172,
and resemble those of A. calva rather than
those of A. fra<s,osa or A. uintacnsis. The
relative munber of teeth and their extent
on the vomer cannot be discerned.
As in A. calva and A. uintacnsis, the den-
tary in A. scutata lacks the dorsal shelf
adjacent to the lingual border of the alveo-
lar ridge seen in A. fra^osa (Fig. 18). The
bone is very thick, especially toward the
mid-lingual surface, where the dorsal and
ventral halves meet to form the Meckelian
groove. As in A. uintacnsis the upper wall
of this groove is primarily formed bv the
thickness of bone in dorsal half of the lin-
gual surface; the ventral half is barely
overlain by the dorsal half. The first coro-
noid does not extend past the Meckelian
groove, and bears sharp conical teeth, as in
A. calva (Fig. 18). The anterior half of the
dentary is more incurved than in A. uintacn-
sis, but not to the extent that it is in A.
calva. The anterior width of the dentary
also resembles that of A. calva and A. uin-
tacnsis in that it is evenly tapered almost to
the symphyseal edge. The angular and sur-
angular are similar to comparable bones in
A. calva, except that they, like the dentary,
are more extensively ossified. The mandi-
ble/head-length ratio in A. scutata is well
within this ratio range for A. calva (Table
7). The prearticular is not prc\served.
The preoperculuin is similar to that of
A. uintacnsis, being narrower dorsally dian
ventrally, radier than having both halves
rc^latively equal in widdi, as in A. calva.
The operculum resembles that of A. calva
in morphology and opercuhun-depth/oper-
culum-length (Table 7). The suboperculum
and interoperculum resemble those of A.
calva in general morphology, but the suture
between them is longer anteroposteriorly.
The branchiostegal rays are squared off
distally, as in A. calva.
Post-cranial skeleton. The supracleithnmi
and metacleithriun are not preserved. The
only part of the cleithrum available for
study is the ventroposterior process in
UMMP V-57431 which in A. calva is the
only area of this bone that is visible ex-
ternally. This region of the cleithrum in
A. scutata is heavily sculptured (Fig. 21),
and as in A. uintacnsis and A. fra^osa, this
d(Tmal ornauKMitation extends to the edge
of the bone. In A. calva, diis d(>rmal struc-
ture is limited to the cent(>r and dorsal re-
gion of this part of the cleithrum.
Th(> vertebral column of A. scutata re-
sembles that of A. calva both in number of
ccMitra (Table 9) and in general morphol-
ogy of the centra. The head /standard-
length proportion (0.312) is greater than in
A. calva (0.271 ), but less than in A. uintaen-
70 Bulletin Museuui of Comparative Zoology, Vol. 146, No. 1
sis (0.322). The insertion of peetoral fin/
standard-length and insertion of anal fin/
standard-length ratios nrv both within the
ranges of A. calva, althongh the latter pro-
portion for A. scutata is somewhat greater
than the mean for A. calva ( F'ig. 31 ).
Discussion
In the same paper as his deseription of
Aryiia scutata. Cope (1875: 3) described
Amia clictyocepliala, also from the Florissant
Formation. A. dictyocephala was distin-
guished from A. scutata by having 10 to 12
supravertebral scale rows, and 35 vertebrae
between the anterior dorsal fin pterygio-
phore and the posterior anal fin ptervgio-
phore (USNM 3992 AMNH 2670). "lie
further characterized this form from a skull
(AMNH 2S02) that "possesses twelve
branchiostegal rays, and a relatively smaller
orbit than in Amia calva." A re-examination
of these specimens in the previous section
on meristics showed that Cope's supra-
vertebral scale row count was in error, and
there is no perceptible difference in this
feature between Recent and fossil Amia
species (Table 8). In A. calva, the range
for the number of centra between the in-
sertion of the dorsal fin and the terminus
of the base of the anal fin is 33-37. In the
type specimen of A. dictyocephala (USNM
3992) the number of centra is 35, and the
mean number in specimens of A. scutata is
36; there is clearly no way that this fcnitiue
can be used to distinguish A. dictyocephala
from A. scutata and A. calva. Cope, on the
basis of AMNH 2802, thought that an orbit
in A. dictyocepJiala was smaller than one in
A. calva, but the small size was due largely
to the constriction of the orbit that resulted
from crushing of the dcrmosphenotic and
upward displacement of infraorbital 5. The
characters that Cope used to differentiate
A. dictyocephaki from A. scutata are un-
diagnostie, and my studies of the specimens
show no morphological or meristie differ-
ence; A. dictyocepJiala is here considered
to be a synonym of A. scutata.
Lambe (1908: 12-13) described Amia
exilis from a single basioccipital (NMC
6201 ) and two mid-trunk \ c>rtebrae ( NMC
6200, 6205) from the Farly Oligocene
Cypress Hills Formation of Saskatchewan.
The temporal occurrence of these elements
is equivalent to that of A. scutata. Lambe's
description of the basioccipital conforms to
that of A. scutata in being more extensively
o.ssified than in A. calva. His diagnosis of
the two centra is founded on height/ width
proportions, ehordal foramen position, basa-
pophyseal angle, and configuration of nc>ural
facets. Because A. scutata resembles A.
calva in vertebral morphology, the charac-
ters that Lambe uses to distinguish A. exilis
are undiagnostic; I therefore consider A.
exilis iis a synonym of A. scutata.
Amia cf. scutata
Hypodi^m. Miocene. Pawnee Creek
Formation, Colorado: UCMP 38222, nearly
complete cranial roof, infraorbitals 4 and 5,
nearly complete anterior portion of palate,
two branchiostegal rays, maxillae, and right
dentary.
Description
The general morphology of the cranial
roof resembles both A. scutata and A. calva
in parietal/ frontal ratio (Table 7), rectan-
gular parietals, and shape of dermosphen-
otic and nasal (Fig. 29). The extrascapular
more closely resembles that of A. scutata in
its greater width and less pronoimced distal
posterior lappets. The pterotic also resem-
bles that in A. scutata in its being narrower
anteriorly than posteriorly, and in bordering
die frontal laterally rather than posteriorly.
The size and depth of the orbital excavation
is intennediate between that of A. scutata
and A. calva (Fig. 28). The maxilla is
similar to that of A. calva, being less robust
posteriorly than that of A. scutata. The
branchiostegal rays are squared off distally,
as are tliose of both A. calva and A. scutata.
Infraorbital 4, although posteroventrally in-
complete, is clearly closer to that of A. scu-
tata than A. calva in being relatively larger
than infraorbital 5, and in the posterodorsal
corner being squared off rather than acute
as in A. calva. Infraorbital 5 resembles that
I
Fossil Amiids • Boreske 71
of A. scutata in size relative to infraorbital 4,
the anterior end bcnng narrower than in
A. scutata; this featnre eontributes to lessen-
ing the relative width of the orbit. The
dentary resembles that of A. scutata in
being wider anteriorly than in A. calva; the
dorsal lingual surfaee only slightly overlaps
the ventral lingual siuface as in A. scutata
(Fig. 18); Meckel's groove is thus similar
to that of A. scutata. There is no available
palate in A. scutata for comparison. The
number of vomerine teeth is 18 and 21,
which is bracketed by the range for A.
calva (Estes and Berberian, 1969: 5). As
Estes ( 1964 ) noted for this specimen, these
teetli are sharper and more incurved ex-
ternally than internally; this disparity is
more distinct in this form than in the extant
species. The hyomandibular, entopterygoid,
ectopterygoid, dermopalatine, and pre-
maxilla are poorly preserved, but appear
to resemble these bones in A. calva.
Discussion
Estes (1964: 36) and Estes and Tihen
(1964: 454) referred to this specimen as
Amia sp. (and in error gave the source as
White River Formation). The .specimen
resembles A. scutata in some elements, A.
calva in others, and is intermediate in
several character-states, notably bone thick-
ness and size of orbits. It does, however,
appear to show a stronger resemblance to
A. scutata than to A. calva, particularly in
Fig. 29. Amia cf. scutata DC 38222, Late Miocene, Pawnee Creek Formation, Colorado.
72 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
the morphology of the extrascapiilar, ptcro-
tic, dentary, and infraorbitals 4 and 5, and
I have thus compared it with the fossil
species. Since this is a form that is both
morphologically and temporally interme-
diate between A. scutata and A. calva, it is
difficult to determine whether or not this
specimen actually represents A. scutata or
a later stage of evolution leading to A. calva,
but it is at least of interest in documenting
the slow phyletic development toward A.
calva in mid-Cenozoic time.
Amia cf. calva
Hijpodi^m. Pliocene. Lower Valentine
Formation, Nebraska: UCMP 65851, an-
terior portion of left dentary and a trunk
vertebra; UMMP 521S7, right nasal, ectop-
terygoid fragment, unidentified cranial
fragments; UMMP 421S5, right dentary
fragment. Ogallala Formation, Kansas:
UMMP 55574-55578, three right and two
left dentary fragments; UMMP 55579, in-
complete right cleithrum; UMMP 55583, a
right extrascapular; UMMP 55580, a right
maxilla; UMMP 55585, a left premaxilla;
UMMP 55586, several scales.
Discussion
Smith (1962), and Estes and Tihen
(1964) described as Amia sp. a nasal and
dentary, and cranial fragments from the
Lower Valentine Formation, Nebraska.
Wilson (1968) described as Amia calva
denatry fragments, a premaxilla, a maxilla,
an extrascapular, an incomplete cleithrum,
and several scales from the Ogallala Forma-
tion, Kansas. This Early Pliocene material
resembles A. calva more closely than does
the Miocene A. cf. scutata specimen noted
above; the elements are very lightly ossified
as in the Recent species. The cleithrum is
distinctly A. calva-\ike in its lack of distal
marginal dermal sculpture. The dentary
fragments are also thinly ossified as in A.
calva, but are slightly wider relative to the
dentary in the Recent species, as in the
Miocene form. Temporally, this Pliocene
material is later than the Miocene form and
earlier than A. calva; morphologically, how-
ever, the available elements conform with
A. calva.
Amiidae incertae sedis
Hypodi^m. Cretaceous. Paluxy Forma-
tion, Texas: SMUSMP 62270, dentary frag-
ments, premaxillary fragment, vertebrae,
maxillary fragments, and an unidentified
palatal bone bearing teeth; FMNH 7050,
basioecipital; FMNH 7051, mid-trunk ver-
tebra; FMNH 7052, anterior trunk vertebra
fragment; FMNH 7053-7054, anterior trunk
vertebrae; FMNH 7055, caudal vertebra;
FMNH 7056, small vertebrae; FMNH 7049,
unidentified palatal bone bearing teeth.
Description
The dentaries are fragmentary (Fig. 30);
the only diagnostic features available for
comparison with other amiid forms are
related to the anterior region of the dentary.
The surfaces pits on the exterior side of the
dentary are relatively larger and deeper
than in any species of Amia. The dentaries
lack the dorsal shelf adjacent to the lingual
side of the alveolar ridge seen in A. fragosa.
The coronoid articulation surface descends
directly from the alveolar ridge, as in a Uro-
cles dentary from the Late Jurassic (Pur-
beck) of England (BMNH 48236). The
lingual surface above the Meckelian groove
is relatively short, even more so than in
Amia uintaensis, and the groove itself is
quite wide, more so than in BMNH 48236.
The anterior portions of the dentaries are
relatively straight, rather than incurved as in
Amia fragosa, and are evenly tapered to the
symphyseal edge. The dentary and pre-
maxilla teeth are broken, but in dorsal view
the interior surfaces of the broken teeth are
very even, lacking the serrated outline seen
in other species of Amia. Only the anterior
portion of the premaxilla is present; it bears
nine alveoli, conforming in this respect with
all Am,ia species. The premaxilla, although
incomplete, displays the anterior (ventral)
edge of the large foramen that is character-
istic of Amia. Only part of the anterior
maxilla is present in the specimens avail-
able, and since the more diagnostic aspects
Fossil A muds • Boreske 73
occur posteriorly, it is difficult to determine smaller fragment (SMUSMP 62270) bears
any affinities witli particular specic\s; the pillar-shaped teeth with nipple-like tips, as
anterior portions that are available gener- in the tooth-bearing palatal bones in species
ally conform with those of Amia. The of Amia. Posterior to the spinal arterial
specific bones to which the palatal frag- foramina the basioccipital includes one
ments belong cannot be identified. The fused vertebra. As in Amia fra^osa and
Fig. 30. Amiidae incerfae sedis, Early Cretaceous, Poluxy Formation, Texas: A'-A-, anterior portion of left den-
tary; B, premaxlllary fragment; C, anterior portion of rigfit maxilla; D, unidentified palatal fragment. XO-15
74 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
Amia caJva, the basioccipital has an ovoid
articular surface with no dorsal indentations
between the neural facets. The large verte-
brae are thickly ossified, as in the Creta-
ceous specimens of Amia cf. uintaensis.
The chordal foramina are closed and the
only available large mid-trunk centrum dis-
plays the pronounced triangular outline
characteristic of Amia uintaensis. None of
the large vertebrae display the character-
istic Amia aortal facets; they do, however,
possess neural facets, and the mid-trunk
centra bear basapophyses. The small verte-
brae are also thickly ossified and the
chordal foramina of the trunk vertebrae are
closed. As Traquair (1911: 39) noted for
Amiopsis cloUoi, the lateral sides of the
vertebrae are marked by a number of vari-
able excavations, or "oval fossae" (Fig. 7).
These smaller mid-trunk vertebrae, unlike
the large ones, display both aortal and
neural facets, as well as basapophyses and
lateral oval fossae.
Discussion
Thurmond ( 1969 : 88 ) reported "various
fragments of an undetermined amiid" from
the Paluxy Formation of Texas, which is
the earliest known occurrence of amiids in
North America. He further noted that
amiid material occurred both in freshwater
and marine zones and that a further descrip-
tion of this material would be the subject
of a later study. He was uncertain as
to whether the amiids occurring in the
marine zones were actually marine or were
freshwater forms secondarily deposited in
the marine areas. None of the material can
be referred to Atnia since it displays charac-
teristics of Amia uintaensis, Amia fragosa,
Urocles, and Amiopsis, as noted in the
above description. The vertebrae suggest
the possibility of more than one form: the
large vertebrae are subtriangular and re-
semble Am'a uintaensis in morphology,
with the exception of the lack of aortal
facets on the trunk vertel:)rae. The small
vertebrae are Amia frag,osa-\ike in morphol-
ogy; they possess aortal facets, but also
display lateral oval fossae characteristic of
Amiopsis. In reviewing the European Juras-
sic and Cretaceous Urocles, Lange (1968)
found little morphological justification to
warrant continued generic distinction be-
tween Urocles and species described by
Woodward ( 1916 ) as belonging to Amiop-
sis from the Purbeck Beds near Wevmouth,
Dorset. Lack of knowledge of the skull of
Amiopsis makes it impossible to compare
cranial elements with those of other amiids;
the singular postcranial feature charac-
terizing Amiopsis is the lateral o\'al fossae
of the v^ertebrae. Although Lange suggests
that both Amiopsis and Ainia evolved inde-
pendently from different Urocles species-
groups, it is premature to attempt to do
more than indicate morphological similar-
ities or dissimilarities since the phylogenetic
relationship of Amiopsis with Urocles or
Amia cannot be clearly defined until a
much-needed review of the taxon has been
completed, and until more Amiopsis mate-
rial is made available for study.
The Paluxy material shows resemblances
to two early Aryiia species, Amia uintaensis
and Amia fragosa, as well as to the Late
Mesozoic European amiids, Urocles and
Amiopsis. Whether the Paluxy material
represents one or more forms intermediate
between Atnia and Urocles (or Amiopsis)
or whether it belongs to some other group
of amiids that became extinct before the
end of the Cretaceous cannot be deter-
mined, since taxonomic evaluation of this
material is limited by the lack of articulated
specimens.
SPECIMENS REMOVED FROM
THE AMIIDAE
Miller (1968: 468-470, pi. 1, figs. 1, 3,
7-9) questionably identified as Protamia
sp. one large (FHKSCM 13025) and three
small centra (FHKSCM 13024-9) recovered
from a channel sandstone cut into the Up-
per Cretaceous Black Creek Formation,
Phoebus Landing, North Carolina. Since
all known Ainia are freshwater forms and
since these centra were associated with vari-
ous marine vertebrates. Miller (1968: 467)
concluded that the channel sandstone con-
Fossil Amiids • Boreske
75
tained a mixed fauna, "the ehannel sand-
stone formed in an estuarine or tidal en-
vironment."
My studies indicate that these specimens
are not amiid. The smaller vertebrae are
horizontally ovoid. A. uintaensis trunk cen-
tra have concavities between the basa-
pophyses and aortal facets ( Fig. 26, A-C ) .
A thin section (Fig. 26, G) through the
articular surface of one of the Nortii Caro-
lina specimens (FHKSCM 13924-9) has a
radial structure resembling that of S(juatimi
and other sharks (Hasse, 1882, tables 17-
18). All layers are laminated parallel to the
exterior surface and are crossed by various
perpendicular vascular foramina. Their
articular surfaces are slightly concave, while
those of Ainia are markedly so. Each of the
small vertebrae bear horizontal basapoph-
yses as in Recent Sqiialus, and are best
referred to the elasmobranchs.
The large vertebra is a cetacean caudal
(Fig. 26, H), possibly belonging to the
Cetotheriidae (Clayton Ray, 1971, personal
communication). The centrum is ovoid,
with very slightly concave articular surfaces,
and lacks a chordal foramen, as well as
ventral facets. The dorsal facets for the
accommodation of metapophyses are well
defined. Since this centrum is from a ma-
rine mammal, it is more probably from the
Miocene (Calvert Formation?) than from
the Cretaceous Black Creek Formation.
Eastman (1899) described Amiopsis dar-
toni from a partial opercular series, pectoral
fin, and associated cycloid scales from the
Late Jurassic marine Sundance Formation,
South Dakota. Eastman felt that the many
"stout ribs" associated with the pectoral fin
suggested a well-ossified Aniia-hkc verte-
bral column and the semicircular operculum
conformed with that of A. ctilvu. Since the
scalers are covered supt^rficially with ganoine
and appear elliptical, Eastman placed this
form among the Amiidae. He allocated the
generic name, Amiopsis, on a temporal
basis. According to Bobb Schaeffer, (1971,
personal communication) the holotype
(USNM 4792) and the paratypes (MCZ
9696, USNM 4793) are to be tentatively
referred to the Leptolepidae on the basis
of morphology of opercular series and pec-
toral fin lepidotrichia. Schaeffer is currently
studying the Late Jurassic North American
fishes and is including a more extensive
discussion of this material in his review.
SUMMARY AND CONCLUSIONS
This survey of the osteology, morpho-
metries, and meristics of the North Amer-
ican fossil amiids indicates that the extant
and fossil forms fall into foiu- groups
worthy of specific status: ( 1) Amia fragosa,
(2) A. uintaensis, (3) A. scutata, and (4)
A. colvci. All these forms, excepting A.
fragosa, have somewhat elongated bodies
(approximately 85 centra) and shai-p,
conical coronoid and palatal teeth. Al-
though the coronoid and palatal teeth of
A. uintaensis are more sharply curved in-
wardly, the teeth are closer in morphology
to those of A. scutata and A. calva than
to the styliform teeth of A. fragosa. A.
uintaensis, A. fragosa, and A. scutata all
have a larger infraorbital 4 than infraorbital
5, greater degree of ossification of cranial
elements, deeper orbital notch in the frontal,
greater head/ standard-length, and generally
larger parietal /frontal ratio. These charac-
ter-states clearly set the fossil species of
Amia apart from the Recent A. calva.
Articulated specimens have yielded more
information on the osteology of A. fragosa.
A. fragosa is a short-bodied form (approxi-
mately 65 centra) with a smaller number of
caudal lepidotiichia than in the other
species of Amia, styliform palatal and coro-
noid teeth, deeper orbital excavation in the
frontals, square parietals, and a short box-
like skull having relatively short mandibles
diat occupy about half the head-length. The
styliform crushing palatal teetlr of A. fragosa
suggest a durophagous habit, rather than
the more predaceous habit indicated by the
sharp palatal teeth of A. uintaensis, A.
scutata, and A. calva. Although it is known
that A. calva includes molluscs and crusta-
ceans in its diet, perhaps A. fragosa was
more exclusively adapted for shell crushing
than the Recent species.
76 Bulletiti Museuin of Comparative Zoologtj, Vol. 146, No. 1
Fig. 31. Skull and body structure of A, Amia calva; B, A. scufafa; C, A. uinfaensis; and D, A. fragosa.
Fossil Amiids • Boreske 77
PLEISTOCENE
PLIOCENE
POST-BLANCAN
BLANCAN-
HEMPHILLIAN
CLARENOONIAN
FOSSIL LAKE BEOS
(lOAHO FM.)
WAKEENEY It.
(OGALLALA FM.)
LOWER VALENTINE FM.
BARSTOVIAN
MIOCENE
HEMINGFORDIAN
ARIKAREEAN
EUBANKS l.f.
(PAWNEE CREEK FM.)
TURTLE BUTTE FM.
WHITNEYAN
OLIGOCENE
ORELLAN
CHAORONIAN
RUBY PAPER SHALE
(PASSAMARl FM.)
GRANT HORSE PRAIRIE SHALE
FLORISSANT FM.
ORELLA MEMBER
(BRULE FM.)
CHADRON FM.
CYPRESS HILLS FM.
OUCHESNEAN
EOCENE
UINTAN
BRIDGERIAN
WASATCHIAN
CLARNO FM.
HORSEFLY RIVER BEDS
UINTA FM.
WASHAKIE FM.
BRIDGER FM.
WIND RIVER FM.
FOSSIL LAKE BEOS
(GREEN RIVER FM.)
6OLOEN VALLEY FM.
WASATCH FM.
GRAYBULL BEDS
(WILLWOOO FM.)
CLARKFORKIAN
TIFFANIAN
PALEOCENE
TORREJONIAN
PUERCAN
MAASTRICHTIAN
CRETACEOUS
CAMPANIAN
ALBIAN
BEAR CREEK l.f,
(FORT UNION FM.)
SILVER COULEE l.f.
(FORT UNION FM.)
MELVILLE FM.
SAUNDERS CREEK l.f.
(PASKAPOO FM.)
CEDAR POINT QUARRY l.f.
(FORT UNION FM )
MEDICINE ROCKS l.f.
(TONGUE RIVER FM.)
ROCK BENCH l.f.
(FORT UNION FM4
TULLOCK FM.
MANTUA If.
(FORT UNION FM.)
HELL CREEK FM.
LANCE FM.
OJO ALAMO FM.
AGUJA FM.
EDMONTON FM.
JUDITH RIVER FM.
"MESAVERDE" FM.
OLDMAN FM.
BUTLER FARM l.f.
(PALUXY FM)
7X
Table 18. Major deposits containing remains of Amia in the
WESTERN interior OF THE UnITEU StATES AND CaNAUA
78 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
Seven genera and twenty-three amiid
species (Table 19) have been described in
the literature. Estes (1964) synonymized
Stylomyleoclon lacus with Kindleia fra^osa,
and Estes and Berberian ( 1969 ) referred
the genus Kindleia to Amia, thereby con-
firming the suggestion of Janot (1969).
Paramiatus p,iirleyi (Romer and Fryxell,
1928) is unquestionably a synonym of A.
fragosa. Regardless of possible synonymy
with European taxa, the sti'atigraphic
range of A. fragosa is remarkably long,
extending as it does from the Late Creta-
ceous through the Middle Eocene. Al-
though A. fragosa is better known than the
other fossil species, and was extensively
described by Estes (1964), O'Brien (1969),
and Estes and Berberian (1969), its phylo-
genetic relationship to them and to A. calva
could not be understood without compara-
tive information on both the other fossil
forms and A. calva (Fig. 32).
A. newherrianus and A. depressiis
(Marsh, 1871), and A. gracilis (Leidy,
1873a), described from undiagnostic ver-
tebral characters, are considered here as
nomina duhia.
A. iiintaensis is a form having a relatively
greater body-length than the other species
of Amia. It has approximately the same
total number of vertebrae as A. calva and
A. scutata, but the arrangement of the
coliunn varies meristically from them. Its
head is more elongated tlian that of the
other forms, with the jaws occupying over
two-thirds of the head-length. The vouier-
ine teeth are sharp (as are the palatal and
coronoid teeth), as they are in A. scutata
and A. calva, but are more than twice as
numerous as in these later forms. The pres-
ent study confirms the opinions of Romer
and Fryxell ( 1928), Estes ( 1964), and Estes
and Berberian (1969) that the differences
between Amia and Protamia, Hypamia, and
Pappichthys are insufficient for the recogni-
tion of any of the latter as genera distinct
from Amia. Hypamia elegans (Leidy,
1873a) is considered a m)men duhium, be-
ing based on vertebral characters that can-
not be distinguished from those of the
other species. Protamia media (Leidy,
1873a), Pappichthys symphysis, P. corsonii,
P. medius, P. plicatus, P. sclerops, P. laevis
(all described by Cope, 1873), as well as
Atnia macrospondyla and A. whiteavesiana
(Cope, 1891), are all considered here as
synonyms of A. uintaensis; they were based
on undiagnostic vertebral characters and
morphology of the skull elements. Material
of large amiids from the Late Cretaceous
Lance and Hell Creek formations is referred
to A. cf. uintaensis, since the material differs
only in minor respects from the Paleocene
and Eocene specimens. It cannot be deter-
mined whether this material represents ac-
tual populations of A. uintaensis or an
earlier stage of its evolution. The strati-
graphic range of A. uintaensis extends from
the Paleocene to the Early Oligocene.
A. scutata, an Early to Middle Oligocene
long-bodied form, shares cranial characters
with both A. uintaensis and A. calva. Al-
though it has closer morphometric and
meristic affinities to the Recent form, it is
structually and temporally intermediate be-
tween A. uintaensis and A. calva; it resem-
bles the more primitive A. uintaensis in the
moi-phology of Meckel's groove and coro-
noid articulation surface of the dentary,
greater ossification, and in having an elon-
gated skull with a greater head/ standard-
length than in A. calva. A. dictyocephala
(Cope, 1875) is considered a synonym of
A. scutata; it was based on undiagnostic
meristic characters. In the evolutionarv con-
tinuum, A. scutata appears to be an inter-
mediate stage between A. uintaensis and
A. calva (Fig. 32). A more direct line of
evolution exists between A. scutata and A.
calva; this is supported by Miocene and
Pliocene amiid material that displays cra-
nial elements closely transitional between
the two species. Thus the Recent species of
A. calva had begun at least by the begin-
ning of the Pliocene, and A. calva was ap-
parently distinct from A. scutata by that
time. It appears that A. fragosa represents
an amiid population that survived until the
Middle or Late Eocene and had no phylo-
genetic affinities with the modern form be-
yond this time.
I
Fossil Amiids • Boreske 79
/Im/'o colva
RECENT
PLEISTOCENE
POST-BLANCAN
Rl ANT AM
HEMPHILLIAN
PLIOCENE
CLARENDONIAN
Ami a cf. calva
BARSTOVIAN
A mi a cf . scut at a
MIOCENE
HEMINGFORDIAN
ARIKAREEAN
WHITNEYAN
OLIGOCENE
ORELLAN
CHADRONIAN
Ami a scuta ta
- i
DUCHESNEAN
, J
EOCENE
UINTAN
BRIDGERIAN
\
y
WASATCHIAN
^ragosa Amia uintaensis
CLARKFORKIAN
'V"
/
PALEOCENE
TIFFANIAN
TORREJONIAN
PUERCAN
\
\ / -
MAASTRICHTIAN
\ Amia ct uintaensis
CRETACEOUS
CAMPANIAN
\ ;
ALBIAN Al
niidae
mcertae sedi
\
s
Fig. 32. Suggested phylogenetic relationships within the genus Am\a.
80
Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
In the North American fossil record, fossil
remains unquestionably those of the family
Amiidae first occur in tlic Lower Cretaceous
(Albian) sediments of Texas. However,
none of the material can be referred to any
known species of Araia; it displays charac-
ter-states resembling those of Amia uintaen-
sis and Amia frcifi^osa, as well as the Euro-
pean Urocles. Some of the vertebrae re-
semble those of Amiopsis. The Paluxy mate-
rial may represent either one or more forms
transitional between Amia and the Late
Mesozoic European Urocles (or Amiopsis) ,
or an as yet undescribed line. The body-
length of Amia fra^osa appears to be a
primitive feature derived from the earlier
amiids Urocles, Siruimia, Ikechaoam,ia, and
Amiopsis. Despite thc>ir different vertebral
columns, Amia frafi^osa and A. uintaensis
show similar morphology of the cranial
elements, but the nature of the probable
common origin of these forms is still uncer-
tain in the absence of a more complete fossil
record.
Remains of amiids referable to or close to
Amia fragosa and A. uintaensis have been
described from the Paleocene, Eocene, and
Oligocene of Europe, and the Eocene of
Asia. Additional but still not definitive evi-
dence supports Estes' ( 1964 ) and Estes and
Berberian's (1969) suggested synonymy of
A. russelU (Late Paleocene, France), A.
kehreri (Middle Eocene, Germany), and
A. munieri (Early Oligocene, France) with
A. fragosa. Pseudamia Jieintzi (Eocene,
Spitzbergen ) and A. valenciennesi ( Eocene,
France) are also possible synonyms of A.
fragosa. A. valenciennesi is the oldest name
and would take precedence over A. fragosa.
Cranial similarities confirm the synonymy of
A. rohusta (Late Paleocene, France) with
A. uintaensis.
European and North American fossil
Amia occurred in freshwater deposits and
apparently occupied a habitat much like
that of the Recent species. According to
Westoll (1965: 19-20) the distribution of
freshwater vertebrates is a useful indica-
tion of "direct continental communication,"
Table 19. Amiid genera and species of various
AUTHORS discussed IN TEXT IN RELATION TO THE
REVISED lAXONOMY
Atnia calva
Kindlcia fragosa
Stijlomijleodon lactis
Amia fragosa
Paramiattts f^tirlcyi
Amia scutata
A7nia dictyocepliala
Aitiia cxilis
Protamia uintaensis
Protamia media
PappicJi th ys m edius
PappicJithys pJicatus
Pappichthys sclerops
Pappichthys lacvis
PappicJithys symphysis
PappicJithys corsonii
Atnia loJiiteavcsiana
Amia macrospondyla
Amia depressus
Amia newJwrriamis
Amia gracilis
Hypamia elcf^ans
Arnia sehvyniana
. . Amia calva
Amia fragosa
.Amia sctitata
Amia uintaensis
nomina dubia
.nomen nudum
since ". . . descendents of a common stock on
different modern continents must have used
essentially a terrestial route." The present
study further amplifies similarities in the
Paleocene and Early Eocene amiid fossil
record of North America and Europe. This
distribution of amiids adds to the similarity
of assemblages of Paleocene and Early
Eocene lower vertebrates (Estes et al.,
1967) and mammals (McKenna, 1972) on
the two continents. The occurrence of
Pseudamia Iwintzi in the Eocene deposits
of Spitzbergen may be additional evidence
for the existence of the De Geer migration
route (bridging Europe, Spitzbergen, and
North America during the Paleocene and
until the close of Sparnacian time), espe-
cially if suggested relationship to A. fragosa
could be demonstrated. The Asian form A.
mongoliensis resembles A. uintaensis in
minor respects but is sufficiently distinct in
itself to be maintained as a separate species.
Fossil Amiids • Boreske 8i
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5 cm
afab.
B
.J<Vj,
Plate 1. A, "Paramiatus gurley'i" FMNH 2201, Early Eocene, Green River Formation, Wyoming; B, Amia iragosa
MCZ 5347, Early Eocene, Green River Formation, Wyoming.
Fossil Amiids • Boreske 85
Plate 2. Amia kebreri BMNH P33480, collected by Walter Kijhne in 1951 from Middle Eocene deposits at Messel
bei Darmstadt.
86 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1
Plate 3. Amia uinfaensis PU 13865, Early Eocene, Green River Formation, Wyoming.
Fossil Amiids • Bnrrslr
?^" ■_;.■■•«;>;
m
.: '-^
-■
1 y
•^fe%i
' •
■•■-Tf^
^
•'i.
- . i
Plate 4. Amio scu/ofa. Middle Oligocene, Florissant Formation, Colorado: A, counterpart YPM 6243; B, counter-
part USNM 4087; C, PU 10172.
I
i
us ISSN 0O27.4100
Bulletin OF THE
Museum of
Comparative
Zoology
An Analysis of Variation in the Hispaniolan
Giant Anole, Anolis ricordi Dumeril
and Bibron
ALBERT SCHWARTZ
HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSETTS, U.S.A.
VOLUME 146, NUMBER 2
19 APRIL 1974
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OccAsioNAL Papers on Mollusks, 1945-
Other Publications.
Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine.
Reprint
Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of
Insects.
Creighton, W. S., 1950. The Ants of North America. Reprint.
Lyman, C. P., and A. R. Dawe (eds.), 1960. Symposium on Natural Mam-
malian Hibernation.
Peters' Check-list of Birds of the World, vols. 2-7, 9, 10, 12-15.
Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echinoderms.
Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredinidae
(MoUusca: Bivalvia).
Whittington, H. B., and W. D. I. Rolfe (eds.), 1963. Phylogeny and Evolu-
tion of Crustacea.
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Authors preparing manuscripts for the Bulletin of the Museum of Comparative
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© The President and Fellows of Harvard College 1974
AN ANALYSIS OF VARIATION IN THE HISPANIOLAN GIANT
ANOLE, ANOLIS RICORDI DUMERIL AND BIBRON
ALBERT SCHWARTZ'
Abstract. The nominal Hispaniolan species of
giant anole, Anolls ricordi, is considered to be in
actuality composed of three distinct allopatric spe-
cies: A. ricordi, A. barahonae, and A. baleatus.
Subspecies of all three species are described, but
only A. baleatus is well represented in collections.
A theoretical history of this species complex upon
Hispaniola is presented.
The Hispaniolan giant anole, Anolis ri-
cordi Dumeril and Bibron, 1837, has been
known to science for more than a century;
yet only in the hist 35 years has it become
evident that this species is not homoge-
neous in its characteristics throughout Haiti
and tlie Republica Dominicana. The spe-
cies was first named (as Anolis ricordii)
from Santo Domingo, as the entire island
was known at that historical period, but
specimens seem to have been rare in col-
lections thereafter. Schmidt (1921: 10) re-
ported four A. ricordi from two Dominican
localities. Cochran (1941: 133) Hsted 24
specimens (all but one of which were in
the National Museum of Natural History)
from 11 localities. Mertens (1939: 68-70)
studied 17 specimens in European collec-
tions and was the first to recognize that
there were two readily distinguishable pop-
ulations that he considered subspecies: A.
r. ricordi in Haiti, and A. r. baleatus Cope
in the Republica Dominicana. Williams
( 1962 ) reviewed the species in more detail
and examined 90 specimens. For this suite
of anoles, he described A. r. barahonae
\
1 Miami-Dade Community College, Miami,
Florida 33167.
Bull. Mus. Co
from tlie Sierra de Baoruco in the south-
western Republica Dominicana. Still later,
Williams (1965) studied an additional 80
specimens and named A. r. leberi from
Camp Perrin on the extreme distal portion
of the Haitian Tiburon Peninsula. Thus,
with increasing quantities of material from
more diverse localities, our knowledge of
the distribution and variation in this species
has increased accordingly.
A great many problems remain, however,
when one deals in detail with the variation
in A. ricordi. Williams (1962, 1965)
pointed out that records of the species were
of such a scattered nature (especially on
the Tiburon Peninsula but also elsewhere
on the island) that intergrades between
several of the subspecies remained unknown
and also that there were no specimens
available from large areas between named
populations. Williams and Rand (1969), in
their excellent summary of the geographic
differentiation in all species of Hispaniolan
anoles, pointed out (p. 15) that Anolis ri-
cordi was composed of "several described
subspecies, some of which are sharply
enough distinct to raise the question of pos-
sible species status." This is most especially
true of the taxa ricordi, baleatus, and bara-
honae, all of which are extremely well
characterized by both pigmental and struc-
tural details, but all of which occupy areas
(extensive in the cases of ricordi and ba-
leatus) without known intergradation be-
tween them or without close geographic
approximation. Thus, the closest ap-
mp. Zool., 146(2): 89-146, April, 1974 89
90
Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
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HiSPANiOLAN Giant Angle • Schwartz 91
proaclu's of the ranges of ricordi and 1)0- material upon whieh A. r. leheri was based.
leatits (Copey and Peiia, both in the Our unique experience at Camp Perrin —
Republica Dominieana) are separated by namely, of many A. ricordi received from
some 115 kilometers airline. The subspe- local Haitians — showed that the species
cies harahomie and ricordi (Sierra de Bao- may not be necessarily rare. However, as
rueo and associated eastern coastal areas in A. equestris in Cuba, the cryptic greens
in the Republica Dominieana, and Saltrou of A. ricordi render the species virtually in-
in Haiti) are known from localities sepa- visible during the day except to all but the
rated by about 115 kilometers, and bara- most experienced observer. In 1963, Rich-
honae and balcatus by a gap of about 115 ard Thomas discovered that A. ricordi
kilometers (between the Sierra de Baoruco might be seemed at night, since individ-
and near Villa Altagracia, both in the Re- uals sleep quite exposed in a variety of ar-
publica Dominieana). boreal situations and are very conspicuous.
Schwartz and Garrido (1972) recently Thus, with the knowledge that the agroma
showed that the Cuban giant anole, Anolis (as the species is known in Haitian Creole)
equestris Merrem, is, in fact, a complex of or the saltacocote ( as the species is known
five species; they also suggested (p. 71) in Dominican Spanish) might be common
that, as Williams and Rand had pointed and thus easily secured by native collectors,
out, there was a good possibility that the and that individuals might be readily se-
Hispaniolan Anolis ricordi in time might cured at night while they slept, I had as
likewise be shown to be a complex of spe- one of my objectives to secure as many A.
cies. It is the purpose of the present paper ricordi as possible in order to clarify the
to discuss the variation in A. ricordi, based status of the named subspecies and in an
upon the examination of 403 specimens attempt to narrow the geographic gaps that
from a broad selection of geographic local- seemed to exist between ricordi, baleatus,
ities throughout Hispaniola. Despite my and barahonae. As more material accumu-
ha\'ing studied far more material than any lated, we were successful in the latter at-
previous investigator, there still remain tempt, but the range of variation in newly
many problems that cry out for solution, acquired material showed that the situation
As Schwartz and Garrido also pointed out was more complex than was supposed. In
in their analysis of Anolis equestris, the addition to specimens in the Albert
present paper in no way should be consid- Schwartz Field Series (ASFS), collected
ered as the final word on A. ricordi; rather by myself and field assistants, I have ex-
it is an attempt to comment in detail upon amined material in the American Museum
the known variation and distribution of of Natural History (AMNH), the Museum
this species in Hispaniola which may serve of Comparative Zoology (MCZ), and the
as a stepping stone for further work upon National Museum of Natural History
the species. ( USNM). For the loans of specimens I am
Between 1962 and 1971, I and my asso- grateful to Richard G. Zweifel, George W.
ciates collected extensively in both Haiti Foley, Ernest E. Williams, and George R.
and the Republica Dominieana. Latterly, Zug. In all of these collections there are
between 1968 and 1971, my work in His- other specimens that I have deliberately
paniola has been under the sponsorship of not elected to study, since many of them
two National Science Foundation grants, are from localities that are now well repre-
GB-7977 and B-023603. Specimens of Ano- sented by more recently collected lizards
lis ricordi collected in 1962-63 were avail- or that have poor locality data. Specimens
able to Williams and were reported upon in the collection of the Museum of Com-
by him (1965); in fact, the long series of parative Zoology have been collected un-
A. ricordi from Camp Perrin, Haiti, secured der NSF grant B-019(S01X and previous
for me by native collectors in 1962, was the grants to Dr. Williams. Most of the re-
92 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
cently taken ASFS A. ricordi have detailed
descriptions of color and pattern in life,
but, as in all such endeavors that span sev-
eral years, it is unfortimate that all details
of color and pattern have not been consis-
tently recorded as time has passed. Like-
wise, there are no color or pattern data on
most old specimens; thus, I feel less secure
in dealing with these older specimens or
those collected by others than myself and
parties than I am with those in the ASFS
which are carefully documented. However,
specimens from other localities must in
some way be dealt with, and I have done
so as carefully as possible, considering de-
tails of geography and what is known about
specimens of A. ricordi from adjacent local-
ities.
I wish to acknowledge with enthusiasm
the efforts on my behalf in the field of the
following men, without whose efforts the
quantity of A. ricordi presently available
to me would be far less: Jeffrey R. Buffett,
Carl Butterfield, James R. Dennis, Danny
C. Fowler, Ronald F. Klinikowski, David
C. Leber, James A. Rodgers, Jr., Bruce R.
Sheplan, and Richard Thomas. C. Rhea
Warren has given me a specimen of A. ri-
cordi from northern Haiti. My notes on
coloration and pattern of A. ricordi have
been greatly supplemented during the
present study by the color portraits exe-
cuted in the field by D. C. Leber; one of
these has been reproduced in black-and-
white in Williams ( 1965 ) , but the repro-
duction hardly does justice to the detailed
beauty of all the originals. I have been able
to examine the holotype of Eupristis ba-
leattis Cope through the courtesy of Alice
C. C. Grandison and A. F. Stimpson of the
British Museum (Natural History). Holo-
types and paratypes have been designated
or deposited in the above collections and
in the Caniegie Museum (CM).
THE PROBLEM
Mertens ( 1939 ) was the first to point out
that Haitian and Dominican A. ricordi dif-
fered from each other in one notable char-
acter— the height of the dorsal crest scales.
His figure 41 shows this character ex-
tremely clearly: in nominate ricordi from
Haiti, the nuchal crest scales are low and
inconspicuous, whereas in Dominican ba-
leatiis the nuchal crest scales are long and
attenuate. In addition, Mertens (1939: 69)
characterized ricordi as having 9 to 12
scales between the eyes; males of this sub-
species have one or more sharply defined
black blotches on the sides of the nape, the
occipital area flecked with black, and often
have black longitudinal stripes on the
flanks. On the other hand, baleatus has
from 6 to 8 scales between the eyes, and
males are without any black head, nape, or
lateral markings. Williams (1962) com-
pared these two taxa with barahonae in re-
gard to four characters: height and relative
length of nuchal and dorsal crest scales,
number of snout scales at the level of the
second canthal scale, and body pattern.
Later, when he described A. r. leberi, Wil-
liams (1965) employed these same charac-
ters to differentiate that subspecies.
The differences in these characters be-
tween the four recognized subspecies are
unequivocal: one can differentiate at a
glance between such distinctive animals as
leberi and barahonae or between ricordi
and baleattis, without recourse to micro-
scopic examination. The whole a.spects of
all four taxa are quite distinctive, whether
one is dealing with living or long-preserved
animals. What has been equivocal is the
relationships of these four taxa, since, as I
pointed out previovisly, they have been
known from rather isolated groups of lo-
calities, widely separated from each other.
In only one case (leberi-ricordi) have
specimens been regarded as intergradient
between two subspecies: these intergrades
are from a geographically plausible locality
that itself is widely removed from the two
"parent" populations.
As material has gradually accumulated,
it has become increasingly obvious that the
situation is even more complex than has
been previously recognized. For example,
in 1971, I had occasion to compare long se-
ries of living examples from the Peninsula
HisPANioLAN Giant Anoll. • Schwuiu
de Sainana and the adjacent "mainland" at
Cafio Abajo, and I was at once struck with
the differences between tliese two samples,
both of which have been regarded as ha-
leattis. In this case, the differences are not
particularly subtle but they do involve dif-
ferences in coloration and pattern which
often are evanescent after preservation.
The same statement may be made about A.
ricordi from the region near La Vega and
those from the Cordillera Septentrional. In
1971 I had occasion to collect specimens
from both these regions on two successive
days and thus was able to compare freshly
collected material directly. Again, the dif-
ferences are ones of pattern and color, but
they are so striking that it is misleading to
consider both these populations as being
identical genetically. I could multiply the
above examples but to no purpose; it is ob-
vious, when one sees living A. ricordi in the
field, that there are several populations
presently assigned to haleatus which are
quite distinctive.
On the basis of specimens collected by
Richard Thomas and myself in 1963, Wil-
liams ( 1965 ) reported A. r. ricordi for the
first time from the northwestern Republica
Dominicana in the region near Pepillo Sal-
cedo and Copey in Monte Cristi Province.
He noted, however, that, "Despite the new
collections one embarrassment remains. No
certain intergrades between the two strik-
ingly different forms ricordii and ])aleatiis
are yet known. . . . However, the area in
which intergrades may occur is being nar-
rowed: on the north coast of the Dominican
Republic between Monte Cristi and Santi-
ago and in the center of Hispaniola be-
tween Mirebalais ( MCZ 68479, 69404) and
Santiago. This still leaves a very wide area
of ignorance." Since the above was written,
I have secured specimens of the nominate
subspecies in four other regions: at Re-
stauracion, Dajabon Province, along the
Dominico-Haitian border and about 40 ki-
lometers airline south of the Monte Cristi
localities; on the southern slopes of the Cor-
dillera Central near Juan de Herrera in San
Juan Province; and throughout the Sierra
de Neiba between Hondo Valle and Valle-
juelo in La Lstrelleta and San Juan prov-
inces. These latter two regions (the south-
ern slopes of the Cordillera Central and the
Sierra de Neiba) are separated by the
rather xeric Valle de San Juan. Elsewhere
to the east and north, the Cordillera Cen-
tral harbors A. ricordi with long nuchal
crest scales and without black nape and
head markings in males (i.e., — haleatua),
as at San Jose de Ocoa, La Vega, and the
interior uplands near El Rio, and near Los
Montones on the Rio Bao. The fourth lo-
cality is perhaps the most significant; there
is one subadult male from Santiago Ro-
driguez Province near Los Quemados which
is clearly a ricordi. Of the haleatus locali-
ties, this one is closest to Los Montones, a
distance of 50 kilometers airline. Thus the
gap between ricordi and haleatus in north-
ern Republica Dominicana has been more
than halved, and there still is no genetic in-
fluence of one subspecies upon the other.
To the south, specimens of A. ricordi
from the Sierra Martin Garcia, and Azua
and Peravia provinces likewise narrow the
gap there between haleatus and ricordi on
one hand and between haleatus and hara-
honae on the other. In the former case, the
distance between ricordi and haleatus
(Vallejuelo and Sierra Martin Garcia) is
about 60 kilometers airline, without char-
acter dilution. In the instance of harahonae
and haleatus, only 20 kilometers separate
known localities (Barahona and Sierra
Martin Garcia) for these two taxa: how-
ever, the actual kilometrage is deceiving,
since, lying between these localities, is the
Valle de Neiba and the Bahia de Barahona.
Although this eastern extreme of the Valle
de Neiba is rather mesic and supports (or
did support ) large trees in many areas that
would presumably be suitable for A. ri-
cordi, the break between these two popu-
lations is sharp despite presumably suitable
ecology. I have little doubt that A. ricordi
occurs in this intervening region, and the
interaction of harahonae and haleatus
therein will be most interesting to ascertain.
Even more intriguing is the fact that the
94 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
Sierra Martin Garcia is an extreme eastern
isolate of the Sierra de Neiba, which is
elsewhere occupied by A. r. ricordi. This
small range, which reaches an elevation of
1350 meters, is completely surrounded by
extremely arid desert or low rolling xeric
hills, as well as by the Bahia de Barahona
on its southwestern edges. The heipeto-
fauna of the Sierra Martin Garcia is just
becoming knowti, and it supports a remark-
able fauna, including an endemic species
of Diplog,Iossus (Thomas, 1971) and a new
species of Sphaerodactylus, as well as other
unexpected novelties. Nevertheless, A. ri-
cordi seems to have reached this range
from the northeast (i.e., the southern slopes
of the Cordillera Central), since I regard
the Martin Garcia lizards as identical to
those from Azua and Peravia provinces.
Finally, the geographic relationship of
barohonae and ricordi has been to some ex-
tent clarified. A. r. harahonae has been
known only from the eastern uplands of
the Sierra de Baoruco and from three
southern lowland localities ( Enriquillo,
half-way between Enriquillo and Oviedo,
and Oviedo). Each of the latter localities
is represented by a single specimen. The
Enriquillo and Enriquillo-Oviedo speci-
mens are quite obviously harahonae, but,
as Williams (1965: 4) noted, the specimen
from close to Oviedo is quite different in
style of pattern and color from typical
harahonae. To the west, in Haiti, there has
been but a single specimen from Saltrou
which Williams (1965: 2) considered A. r.
ricordi and which "narrows the geographic
gap between ricordii ricordii and r. hara-
honae; however, it does nothing to narrow
the character gap." Two additional facts
are important. First, in 1971, we secured
a pair of A. ricordi from along the Domin-
ico-Haitian border north of Pedernales;
these individuals, although differing some-
what from typical harahonae and strongly
from the single Oviedo specimen, in no
way show any tendencies toward A. r. ri-
cordi. They are clearly related to hara-
honae, a rather surprising fact since they
are much closer (35 kilometers) to Saltrou
than they are to any harahonae locality (65
kilometers at Enriquillo). Secondly, Wil-
liams has recently received a fine se-
ries of A. ricordi from Source Carroye
near Thiotte; these lizards are quite obvi-
ously not A. r. ricordi but are closer in
many ways to the far-western A. r. leheri.
Thus the situation along the southern Hai-
tian coast between Saltiou and the eastern
coast of the Republica Dominicana at Ovi-
edo and its environs remains a true puzzle.
It seems likely that A. r. ricordi does not
cross the high Massif de la Selle, except
possibly by some circuitous route, and that
A. r. harahonae occurs up to the Dominico-
Haitian border, to within 11 kilometers of
a station ( Source Carroye ) where another
taxon occurs, without character dilution.
Interpretations of all these facts are seri-
ously hampered by the lack of specimens of
A. ricordi from throughout the Haitian Ti-
buron Peninsula. Material from the penin-
sula may be divided into four basic lots: a
short series from the region about Castillon
on the northern slopes of the Massif de la
Hotte near the tip of the peninsula; a very
long series of specimens from Camp Perrin
on the low southern slopes of the Massif de
la Hotte (the type series of A. r. leheri);
a short series from midway along the pen-
insula at Miragoane-Paillant; and a moder-
ate number of specimens from near the
base of the peninsula in the region
of Morne Decayette-Petionville-Port-au-
Prince. The lack of material from such
well-collected areas as Jeremie on the
northwestern coast or Jacmel and Les
Cayes on the southern coast is extremely
puzzling — we simply know nothing about
lowland A. ricordi throughout much of the
Tiburon Peninsula, except for the above
scattered records and the southern coast at
Saltrou near the Dominican border (and
the latter locality is not even on the penin-
sula proper).
To summarize the data from elsewhere
in Haiti and the Republica Dominicana,
there is a huge distributional hole in cen-
tral Haiti, with but two specimens {ri-
cordi) from Mirebalais, widely separated
HisPANiOLAN Giant Anoli
b.:,/Ui i:
from southern ricordi at and near Port-an-
Prince, and then a group of seattered local-
ities along the northern littoral of Haiti
from Port-de-Paix in the west to Terrier
Rouge in the east, and one specimen from
Marmelade in the interior Chaine de Mar-
melade. All these Haitian specimens have
low nuchal crest scales, and males variably
possess some black nape and side markings,
but there are differences between speci-
mens from the various sections which pres-
ently defy analysis, since the material is
too meagre and from too scattered locali-
ties.
The Republica Dominicana fares far bet-
ter as far as detailed coverage is con-
cerned. Aside from the material previously
noted as assigned to ricordi or harahonae,
there are now good series available from
the eastern half of the country, and, al-
though there are certain gaps even within
this region, they are not so appalling as are
those in Haiti. A. ricordi is rarer (or per-
haps less easily encountered) in arid re-
gions, and thus the most striking gaps in
the western half of the Republica Domini-
cana are those involving arid regions on
the one hand or high mountain masses on
the other. The latter situation, especially if
the slopes are pine-clad, does not appear
suitable for A. ricordi. and the species may
be truly absent from the uplands above
4000 feet (1220 meters), the highest eleva-
tion from which the species is known. In
arid regions, A. ricordi appears to be re-
stricted to riverine woods and forests; in
such situations, the species may not be un-
common, but it may require diligence to
secure even one specimen from a particular
region.
The detailed discussion above should
give the reader a background of both the
history and present knowledge of the dis-
tribution of A. ricordi against which the
following accounts can be most logically
followed. One further point is of interest.
A. ricordi is unknown by specimens from
any of the large Hispaniolan satellite is-
lands. In some cases (Isla Beata) the spe-
cies is not expected for a variety of reasons,
but in others (He de la Gonave, lie dc la
Tortuc, Ile-a-Vache) there seems no logical
reason for the absence of A. ricordi, dis-
counting the vagaries of overseas transpoit.
The species does occur on Isla Saona, but
remains uncollected there. Fowler and
Sheplan saw a sleeping A. ricordi on the
northern coast of Isla Saona in December
1971, but, after it had been shot, it was
lost in the undergrowth. The occurrence of
A. ricordi on any Hispaniolan satellite is
noteworthy, and it will be most interesting
to determine the status of the Isla Saona
population.
METHODOLOGY
The series of 403 A. ricordi was divided
into 14 samples on the basis of geography,
as follows: Republica Dominicana: 1) Pe-
ninsula de Samana (54 specimens); 2)
northeastern Republica Dominicana, from
Duarte and eastern La Vega provinces east
to the haitises region in northeastern San
Cristobal Province ( 37 ) ; 3 ) extreme east-
ern Hispaniola, Punta Cana-Juanillo, Boca
de Yuma, La Altagracia Province ( 16 ) ; 4 )
southeastern Republica Dominicana from
Higiiey and Las Lisas, La Altagracia Prov-
ince, west to Santo Domingo and Yamasa,
San Cristobal Province (61); 5) lowlands at
the northern base of the Cordillera Central
at Guaigui, La Vega Province (21); 6)
Cordillera Central at and above elevations
of 2000 feet (18); 7) Cordillera Septentri-
onal and north ( 15 ) ; 8 ) Sierra Martin Gar-
cia and Peravia and Azua provinces (6); 9)
Sierra de Baoruco and associated east coast
of the Peninsula de Barahona (33); 10)
Oviedo, Pedemales Province ( 1 ) ; Haiti:
11) Saltrou and vicinity, Dept. de I'Ouest
( 15 ) ; 12 ) region about Port-au-Prince,
Mirebalais, northern Haitian littoral,
Chaine de Marmelade, and (in the Repu-
blica Dominicana) region about Pepillo Sal-
cedo, Copey, Los Quemados, Restauracion,
and the southern slopes of the Cordillera
Central and the Sierra de Neiba (50); 13)
Camp Perrin and Marceline, Dept. du Sud
(54); and 14) vicinity of Castillon, Dept.
du Sud (6). This division into 14 regional
96
Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
samples was completed after preliminary
examination of the material, scale counts
and detailed review of field notes on color-
ation and pattern were all made. In addi-
tion, two other small lots (eight specimens
from the vicinity of Miragoane, Dept. du
Sud, Haiti, and seven specimens from El
Seibo Province, Republica Dominicana)
were examined separately; in both cases,
these short series indicate intergradient
tendencies between pairs of the 14 major
samples noted above, and they were not in-
cluded with the latter.
The following data were taken on each
specimen:
1 ) Snout-vent length, in millimeters.
2) Number of snout scales across snout
at level of the second canthal scale, reck-
oned from the anterior corner of the orbit.
3) Number of vertical rows of loreal
scales.
4) Minimum number of scales between
supraorbital semicircles.
5) Number of scales between the inter-
parietal scale and the supraorbital semicir-
cles on each side, this datum written as a
fraction ( i.e., 5/5 = five scales in this posi-
tion on each side ) .
6) Number of scale rows between the
subocular scales and the supralabial scales.
7) Number of vertical rows of dorsal
scales in a distance equal to that from the
tip of the snout to the anterior bony wall
of the orbit, this distance measured by ver-
nier calipers, laid off on the back about
three rows below the median dorsal crest
scales, and the number of scales counted
under a binocular dissecting microscope.
8) Number of horizontal rows of dorsal
scales in the snout-eye distance, the scales
counted at midbody. This count was not
taken in most juveniles or on those speci-
mens that were shrunken or poorly pre-
served since, under the latter circumstances,
some smaller ventrolateral or ventral scales
will be included.
9) Number of transverse rows of ventral
scales in the snout-eye distance.
10) Number of lamellae on phalanges II
and III of the fourth toe.
11) Height of the nuchal crest scales,
categorized as very high, high, moderate,
or low.
12) Height of dorsal crest scales, cate-
gorized as high, moderate, or low.
13) Dorsal coloration and pattern of
males and females, separately.
14) Ventral coloration of males and fe-
males, separately.
15) Color of dewlap, in males and fe-
males, separately.
16 ) Color and pattern of chin and throat
in males and females, separately.
17) Color and pattern of upper surface
of head in males and females, separately.
18) Color of eyeskin.
19) Color and pattern of upper surfaces
of hindlimbs.
20) Color and pattern of juveniles and
subadults.
The above characteristics are variously
useful as far as delimiting the nameworthy
populations of A. ricordi. Detailed com-
ments on these characteristics are made be-
low, with especial attention to pitfalls in
their reliability and usage.
1) The snout-vent length of mature in-
dividuals of both sexes is remarkably uni-
form throughout the entire series. Males
are easily distinguished from females at
any age by the presence of one (occasion-
ally two) pairs of enlarged postanal scales.
Many ASFS specimens have the hemipenes
extruded. Males in general reach a larger
snout-vent length than females; the largest
male (ASFS V29284) has a snout-vent
length of 180 and is from sample (4).
whereas the largest female (ASFS V31397)
has a snout-vent length of 151 and is from
sample ( 12 ) . The mean difference in
snout-vent lengths between the two sexes
is about 10 mm in almost all samples with
the exception of maximally sized individ-
uals in both sexes in sample (2) where the
difference is 3 mm, sample (3) where the
difference is 27 mm, sample (4) where the
HisPANioLAN Giant Angle • Schwartz
97
difference is 32 mm, and sample (7) where
both sexes are of the same size. Whether
tliese exceptions to the 10-mm generahza-
tion are meaningful is debatable. At least
samples (2) and (4) include Icmg series
of specimens, and even samples (3) and
(7) include more than ten individuals.
Adults of the two sexes are readily distin-
guished moi-phologically, since males have
a high tail "fin" that is supported by the
bony extensions of the neural spines; this
feature is lacking in females, although they
may have a much reduced caudal crest in
the form of a low ridge. The terminal half
of the tails of many males is crestless; I at
first considered that this was due to break-
age with subsequent regeneration without
regeneration of the tail "fin." Many speci-
mens have this condition, however, without
any obvious change in basic caudal scale
shape and arrangement, and this is the nor-
mal condition in the tails of males. Often
the uncrested portion of the tail is quite
differently colored or patterned than the re-
mainder of the tail; such cases are due to
regeneration.
2) The number of snout scales at the
level of the second canthal has been em-
ployed as a characteristic to separate the
recognized subspecies. Williams ( 1962,
1965) recorded the following variation in
the four subspecies: ricordi, 7-9; Ijaleatus,
2-5; harahonae, 4-6; and leheri, 4-6 (3-6
on map, fig. 2, 1965: 7). It should be re-
called that Williams himself pointed out
that this count alone would not distinguish
all these taxa from each other. Certainly
overlap between haleatus and ricordi is
nonexistent, and between haleatus on one
hand, and harahoiiae or leheri on the other
hand, the overlap is small. Counts on 403
specimens made by myself do not extend
the parameters of snout scales at all: within
the entire lot of specimens, these scales
vary from 2 to 9, just as in Williams's data.
However, the variation within populations
may be much greater than Williams antici-
pated. For instance, in sample (13), the
counts vary between 2 and 7, and in series
(12) between 4 and 8. Most samples have
three or four categories of number of snout
scales. I am in no way implying that this
is an invalid or poor character for differen-
tiation of populations of A. ricordi, how-
ever; it is, rather, an extremely useful one
but requires amplification and interpreta-
tion.
If we examine only those samples (1-8)
which are assigned to haleatus, the amount
of variation in snout-scales is 2-5, exactly
that assigned to this taxon by Williams.
However, within the broad area covered by
haleatus, there are strong modalities of
snout-scales. In samples (1), (4), (5), and
(6), the mode is 2 scales, whereas in sam-
ples (2), (7) and (8), the mode is 4. Only
one sample, (3), has bimodes of 2 and 4
scales. In harahonae (9) the range is 2-
5 (mode 4). In those samples which are
associated with nominate ricordii (samples
11 and 12), leheri (sample 13) and adja-
cent Castillon material (sample 14), the
range is 2-9, thereby showing complete
overlap in range of this count with that of
both haleatus and harahonae. In fact, in
leheri ( 13 ) alone, the range of snout scales
(2-7) almost embraces that for all other
samples and thus the entire species. Mo-
dalities in this complex of samples are 5
(sample 11), 7 (sample 12), 4 (sample 13)
and 6 (sample 14). Sample (7) is nomi-
nate ricordi.
3 ) The number of vertical rows of loreal
scales ranges from 5 to 10. The greatest
variability is in samples (1), (2), (7), and
(12), where the row counts in each case
are 5-9, 6-10, 5-9, and 6-10. Most samples
have four categories of number of loreal
rows. The modes vary as follows: 5 (sam-
ple 11), 6 (samples 8, 9, 13), 7 (samples 1,
3, 4, 5, 6, 7, 12, and 14), and 8 (.sample 2).
4) The minimal number of scales be-
tween the supraorbital semicircles varies
between 1 and 5; no specimen has the semi-
circles in contact. Modes in general are
very strong, and the usual mode is 3 scales
(samples 1-7; sample 8 has a bimode of 2
and 3 scales); these are all samples that
\
98
Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
are assigned to baleatiis. A mode of 3
scales occurs also in samples (11) and
( 13 ) , and of 4 in samples ( 12 ) and ( 14 ) .
5) The number of scales between the in-
terparietal scale and the supraorbital semi-
circles varies between 3/3 and 7/7. Modes
(which in some cases are quite strong and
in others less so) are: 4/4 (samples 1, 5, 6,
8, 9, 11, 13, 14) and 5/5 (samples 2, 4, 7,
12). Sample (3) is peculiar in having the
mode 4/5 (six of 16 lizards), with adjacent
counts of 4/4 (four lizards) and 5/5 (five
lizards). There is high variability in .this
count; it can be assessed in another fashion,
namely, the frequency with which any
scale count (i.e., 4, 5, 6, etc.) occurs within
the sample, regardless of its pairing with
another count on the other side of the head.
With the use of this technique, the fre-
quency of involvement of 4 scales in the
inteiparietal-semicircle contact varies be-
tween 43 percent (sample 1) and 67 per-
cent (sample 8), and of 5 scales between
52 percent (sample 2) and 64 percent
( sample 7 ) . Of the entire lot of specimens,
there is only one occurrence of 7/7 in this
position ( sample 12 ) but many occurrences
of 3/3 (samples 1, 5, 6, 9, 11, 12, 13, a total
of 12 incidences).
6) The number of scale rows between
the subocular scales and the supralabials is
fairly constantly 1, and this is a strong
mode or the exclusive category in all sam-
ples except sample ( 13 ) . In this lot ( which
is the type series, with a few new speci-
mens, of leberi), 48 percent of the lizards
have the suboculars and supralabials in
contact. Elsewhere, contact is absent in
samples (7) and (11). The frequency of
contact varies in all other samples between
3 percent (sample 9) and 17 percent (sam-
ples 6, 8, and 14). These three samples are
widely separated geographically and the
frequency in none of them even approaches
the very high incidence of contact in sam-
ple ( 13 ) . However, it is noteworthy that
samples (13) and (14) are adjacent geo-
graphically.
7), 8), 9) In reference to all counts in-
volving laying out the snout-orbit distance
on the body, I can do no better than once
more to reiterate the cautions previously
expressed ( Schwartz, 1964; Garrido and
Schwartz, 1968; Schwartz and Garrido,
1972) in reference to Anolis equestris and
members of the genus Chamaeleolis. For
these standard-distance counts, old and
poorly preserved, limp, curled, uninjected,
or otherwise distorted specimens are much
less useful and reliable than properly pre-
served, injected, and positioned lizards.
Luckily, by far the largest quantity of A.
ricordi under study are well preserved.
However, I have abandoned counts of hor-
izontal dorsals on young juveniles, even
well-preserved ones, or on any adults
whose condition precluded taking these
counts meaningfully. The juvenile situation
is peculiar in that invariably, despite the
relatively shorter snout of young specimens,
laying out this distance to count horizontal
rows involved including several rows of ex-
tremely lateral and ventral scales, which
are smaller than true dorsals and which
thus tend to increase the horizontal counts.
I have taken vertical dorsals and ventrals
on juveniles, however, and they do not
skew the data. Of the three standard-dis-
tance counts, those of vertical dorsals and
ventrals are much more reliable than are
those of horizontal dorsals.
Vertical dorsal scale rows vary between
12 and 26, with the low count in sample
( 4 ) and the high count in sample ( 12 ) .
Means range from 15.4 (sample 4) to 21.1
(sample 12). These two represent, respec-
tively, lots assignable to baleatus and ri-
cordi, sensu lato. No generalizations of
mean number of vertical dorsals in refer-
ence to samples and geography can be
made, since the range in samples now as-
sociated with baleatus varies between 15.4
and 19.2, with ricordi 17.3 and 21.1, bara-
honae 17.2, and leberi 16.5 and 16.7. Com-
parisons and significance of vertical dorsal
scale counts are shown in Table I.
Number of horizontal dorsal rows ranges
from 13 (sample 1) to 34 (sample 7).
Means vary between 17.1 (sample 11) to
24.6 (sample 12). The latter sample is that
HisPANioLAN Giant Angle • Schwartz
99
Table I.
Taxon
Comparison of number of vertical dorsal scales in popxjtlations
OF giant Hispaniolan angles
M (±2
standard
error of
mean)
.A
50
c
a
c
c
s
2
a.
■2
"a
£
1
-*-
J
c
+
+
+
+
+
+
+
—
+
—
+
—
+
—
+
—
+
+
+
—
+
+
+
—
—
+
+
+
+
+
+
ricordi 50
leheri 54
stihsolanus 15
harahonae 33
samanae 54
caendcolatus 37
litorisdva 16
scelestus 60
multistrtippus 20
sublimis 18
baleatus 15
21.1 ±
16.5 ±
17.3 ±
17.2 ±
16.6 ±
17.1 ±
15.9 ±
15.4 ±
18.6 ±
19.2 ±
.57
.46
.96
.67
.46
.72
.75
.45
1.06
.70
+
+ +
+
+
+
17.5 ± 1.16
A. r. victilus, A. b. alboceUatus, and A. b. fraiidator are not included. A "-)-" in the table indicates that the two
subspecies involved differ significantly ( non-overlap of two standard errors of mean ) ; a "— " indicates no statistical
difference. Note that the mean of A. r. ricordi differs significantly from the means of all other taxa; that of scelestus
differs significantly from those of all other taxa except litorisilva; and that of sublimis differs significantly from those
of all other taxa except multistruppus and baleatus.
of nominate ricordi, the former a peripheral
isolate of baleatus.
Ninnber of transverse rows of vential
scales varies between 15 (samples 1 and
13 ) and 34 ( sample 7 ) . Means range from
20.2 (sample 13) to 2.5.1 (sample 6); sam-
ple ( 13 ) is leheri.
10) The number of lamellae on pha-
langes II and III of the fourth toe varies
between 27 and 39. The variation in any
population is so great that this count is
meaningless as far as differentiation be-
tween any populations of A. ricordi.
11), 12) Williams (1962, 1965) de-
scribed the relative heights and lengths of
both the nuchal and dorsal body crest
scales. Certainly the differences between
baleatus and ricordi, for instance, are so
very obvious on casual examination that
one has no difficulty in ascertaining the
taxon involved. Williams also pointed out
(1962: 4-5) that in some specimens there
is "sometimes a regular alternation of rela-
tively high triangular single scales and
pairs of much lower, more quadrangular
scales," with the result, on some specimens,
of double crest scale rows on the neck; the
net effect of this condition is a rather in-
discriminate grouping or elongate patch of
multiple nuchal crest scales. Although tliis
condition occurs erraticallv, it seems to be
most predominant in specimens from the
Tiburon Peninsula, but it occurs elsewhere
in nominate ricordi and even occasionally
in specimens assigned now to baleatus.
Such a "hypertrophied" nuchal crest condi-
tion does not completely fit any logical geo-
graphical pattern nor is it totally consistent
within any sample, although there are ten-
dencies toward it as noted above. In any
event, it does not obscure the height of the
nuchal crest scales.
In my own analysis, I have used a
slightly different method in recording
height of crest scales. Nuchal scales were
recorded for each specimen as very high,
high, moderate, or low. Such a verbal quan-
tification is not totally satisfactory, since
the investigator's impressions may change
as the study progresses. To avoid this pit-
fall, I re-examined many specimens that
had been studied earlier in the work and
reconfirmed my own early impressions with
my later ones. Body crest scales were re-
corded as high, moderate, and low.
In nuchal crest scales, very high scales
are those which are very elongate, attenu-
ate, almost spinelike ( but of course flexible,
100
Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
not stiff), with the base much shorter than
the height of the scale. High scales are
those which are shorter and less attenuate
than very high scales, but whose height is
still much greater than the base. Moderate
scales are lower and not attenuate, although
they may be pointed, with the height and
base about equal in length. Low scales are
lower than long. The same categories and
interpretations apply to body crest scales,
although no lizard has the body crest scales
so high as the nuchal scales.
Several other points are pertinent. I have
not used this datiun from juvenile and sub-
adult lizards ( all lizards below 100 mm
snout-vent length) since it is obvious that,
regardless of the taxon or sample, all young
A. ricordi have low nuchal and dorsal crest
scales, which, as the lizard matures, be-
come increasingly more specialized until
the adult condition is reached at about 110
mm snout-vent length. Thus young ricordi
and young baleatus, two taxa that are re-
markably distinct in this feature as adults,
are identical in crest development.
Secondly, it might be assumed that (es-
pecially) nuchal crest scales might be bet-
ter developed in adult males than in adult
females; this excessive development might
reasonably be assumed or construed as a
sexually dimorphic character, with hyper-
development in males. Such does not seem
to be the case, and many female baleatus,
for instance, have very high nuchal crest
scales, as high as those of males. In fact,
comparisons of males and females of indi-
vidual samples show that, within each sam-
ple, there is remarkable consistency be-
tween frequencies of the very high, high,
and moderate categories in both sexes.
Thirdly, as one might expect, there is a
sequence of crest scale heights in the nu-
chal-body series. If the nuchal scales are
very high, the body scales are high; if the
nuchal scales are moderate, the body scales
are low, etc. In no case have I recorded a
transition from very high nuchal crest
scales to moderate body scales, for exam-
ple. There is thus a definite correlation be-
tween height of nuchal scales and those of
the body crest.
13) -19) The color and pattern details of
Anolis ricordi throughout its range need
not be gone into in detail at this point. It is
now sufficient to acknowledge that these
lizards show metachrosis varying from
shades of green and green-gray to brown.
The pattern elements, however, are quite
stable, although the hues involved in the
pattern itself may change with changing
base colors. There is little evidence to in-
dicate that a lizard which is, for instance,
blotched in one color phase will become
crossbanded in another. Thus, despite
changes in hues and even in base colors,
patterns remain constant. It is of interest
to note that greens seem to be the colors
that predominate in the wild. The few A.
ricordi that I have seen during the day
have always been green. The many lizards
that I have seen and collected at night
were almost always green, and usually at
their maximum of pattern expression while
they slept. It is this nocturnal assumption
of the green phase coloration that renders
these lizards so very conspicuous at night
while they sleep on exposed branches,
vines, or among the arboreal greenery.
Even in those populations ( Boca de Yuma,
Sierra de Baoruco) in which the greens in-
volved are not bright, the lizards are still
quite obvious at night. It is only rarely
that one encounters a brown A. ricordi at
night. I have notes on only one instance of
this condition, a subadult lizard from the
Cordillera Septentrional.
15) The dewlap coloration in A. ricordi
is variable. In some populations, males
have a pale yellow to gray dewlap, whereas
in others the males have dewlaps that are
peach or vivid orange. It should be noted
that both sexes in A. ricordi have dewlaps
and that the general hue of the female
dewlap is like that of the males, except that
basally it is usually streaked with browns,
dark grays, or charcoal, and the ground
color is less bright than that of males. In
some regions, the female dewlap is quite
differently colored than that of males.
20) Juveniles and subadults present sev-
eral problems that are presently insoluble.
I suspect that much will be revealed once
HisPANioLAN Giant Angle • Schwartz
101
we know the repertory of pattern and color
in yoinig individuals, but these data are
not axailable nx)w. Although there are
many young specimens at hand, they are
inconsistent within populations or even
small samples. The juxenile color is nor-
mally some shade of green (or browns un-
der stress), most often with two to four
pale (cream, whitish, pale gray) cross-
bands. Many small specimens are a uni-
form green without any dorsal markings.
In other juveniles, the dorsum has many
conspicuous crossbands with two shades of
greens (or browns), separated by promi-
nent bands of pale greens (yellow-green,
pea-green), to give a very tigroid lizard;
this condition persists into the adults of one
population, as does the more simply
banded juvenile pattern noted above in
other populations. The juveniles usually
have dark gray to charcoal dewlaps, re-
gardless of their sexes, and often there are
charcoal or white markings on the neck or
aboN'c the forelimb insertion or somewhere
anteriorly. These variants are discussed un-
der each subspecies below, and there is no
need to go into the details here. However,
I do wish to point out that ju\'enile patterns
are more variable than are those of adults,
and that I do not know how to interpret
this situation.
SYSTEMATIC ACCOUNTS
AnoWs ricordi Dumeril and Bibron
Anolis ricordii Dumeril and Bibron, 1837. Erp.
gen., 4: 167.
Type locality. St.-Domingue; holotype.
Museum National d'Histoire Natmelle 1272.
Definition. A giant species of Hispan-
iolan Anolis characterized by the combina-
tion of moderate size (males to 160 mm,
females to 151 mm snout-vent length),
snout scales at level of second canthal
scales variable, between 2 and 9 (modally
4, 5, 6, or 7 by population) but usually 4
to 9 (97 percent), vertical loreal rows 5 to
10 (modes by population 5, 6, and 7),
scales between supraorbital semicircles 2 to
5 (modes 3 or 4 by population), inteipari-
etal scale separated from supraorbital semi-
circles modally by 4 or 5 scales, vertical
dorsal scale's generally small ( 14 to 26 in
standard-distance), ventral scales relatively
large (15 to 32 in standard-distance), nu-
chal crest scales in both sexes moderate to
low, never \'ery high or high, body crest
scales usually low, subocular scales usually
not in contact with supralabials but one
population is remarkably exceptional in
this character; dorsal body coloration and
pattern some shade of green, in some geo-
graphic regions flecked irregularly with
paler and darker green to give a beadwork
effect; male body pattern either of irregular
black to dark brown blotches on the neck,
occipital region of the head, and on sides
(often delimiting two pale longitudinal
bands) or with three longitudinal dark
brown stripes on each side or with dark
saddles and a bluish green flank stripe or
with a powdery pale blue-green lateral
stripe; females usually without dark dorsal
or lateral markings although in some areas
females have a darker brown reticulum,
three pale gray to yellow narrow cross-
bands, longitudinal black lines, or two pur-
ple flank stripes; dewlap in males variable,
from yellowish gray to peach, bright or-
ange, or deep yellow, in females from
peach to deep yellow or dull orange or
even inky brown or inky blue-black, chin
and throat dull yellow, yellow-green, or
pale blue-green in males, eyeskin dark
( charcoal, dark brown ) to light ( pale blue )
in males, charcoal to pale green in females,
and usually with a prominent pale subocu-
lar semicircle clearly delineated.
Distribution. The Tiburon Peninsula in
Haiti, east to the vicinity of Saltron, Dept.
de rOuest, thence north to the northern
Haitian coast from Port-de-Paix east to Ter-
rier Rouge and thence into the Republica
Dominicana east as far as the vicinity of
Los Quemados, Santiago Rodriguez Prov-
ince, south to Restauracion, Dajabon Prov-
ince; also extending from Haiti onto the
southwestern slopes of the Cordillera Cen-
tral in San Juan Province and in the Sierra
de Neiba in La Estrelleta and San Juan
provinces; altitudinal distribution from sea
level to elevations of about 4000 feet ( 1220
meters) south of Castillon, Dept. du Sud,
102 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
Haiti, and to 3500 feet ( 1068 meters ) west
of Marmelade in the Chaine de Marmelade,
Dept. de I'Artibonite, and 3400 feet ( 1037
meters) south of Elias Pifia in the Sierra de
Neiba, La Esti-elleta Province, RepubHca
Dominicana.
Anofis ricordi ricordi Dumeril and Bibron
Type locality. "St.-Domingue"; here re-
stricted to the vicinity of Port-au-Prince,
Dept. de I'Ouest, Haiti.
Definition. A subspecies of A. ricordi
characterized by the combination of mod-
ally 7 snout scales between second canthal
scales, 7 vertical rows of loreal scales, 4
scales between the supraorbital semicircles,
5/5 scales between the interparietal and the
supraorbital semicircles, high number of
vertical dorsal scales (17-26; mean 21.1),
high number of ventral scales ( 19-32; mean
24.7 ) , nuchal crest scales moderate ( rarely )
to low (usually), body crest scales moder-
ate (rarely) to low (usually), subocular
scales usually in contact with supralabial
scales; males usually with some black lat-
eral markings on the neck and thoracic re-
gion, and on the occipital region of the
head, but at times these markings are ab-
sent (see discussion below), females green
and without definite black lateral markings
but at times reticulate with brown, the re-
ticulum delimiting a pair of clear green lat-
eral stripes or with three pale gray to green
vertical narrow bars; dewlap variable, in
males from peach or pale peach to gray or
yellowish gray, and in females from peach
to blue-gray or inky blue or inky black ( see
discussion beyond).
Distribution. Northern Haiti from Port-
de-Paix east to Terrier Rouge and into the
Republica Dominicana as far east as the
vicinity of Los Quemados, Santiago Ro-
driguez Province, and as far south as Re-
stauracion, Dajabon Province, south in Haiti
to the Port-au-Prince region ( Morne De-
cayette, Diquini, Petionville), as well as
east into the Republica Dominicana in the
Sierra de Neiba and the southwestern
slopes of the Cordillera Central in La Es-
trelleta and San Juan provinces.
Discussion. I have little doubt that the
extensive range that I here ascribe to nom-
inate A. ricordi is incorrect. There are sev-
eral very obvious differences in coloration
and pattern between northern and south-
ern specimens of A. r. ricordi; thus the defi-
nition of the subspecies, in order to include
all pattern variants, is necessarily cumber-
some. The problem is presently unresolv-
able since, other than the series from near
Port-au-Prince and the specimens from
northern Haiti, there are huge areas in
Haiti whence specimens remain unknown.
A detailed discussion of the chromatic and
pattern features in the various segments of
A. r. ricordi is given below.
The series of 50 specimens assigned to
the nominate subspecies shows the follow-
ing variation. The largest male (ASFS
V31395) has a snout-vent length of 160,
the largest female (ASFS V31397) 151;
both are from 4.1 mi. NW Juan de Herrera,
San Juan Province, Republica Dominicana.
Snout scales at level of second canthals
vary between 4 and 8; the mode is 7 (22
specimens). The vertical loreal rows vary
between 6 and 10, with a mode of 7 (20
specimens). There are between 3 and 5
scales between the supraorbital semicircles
(mode 4). There are modally 5 scales be-
tween the interparietal and the semicircles;
5 scales are involved in 53 percent of the
combinations; actual counts are 3/3 (1),
4/4 (10), 4/5 (8), 5/5 (17), 5/6 (9), 6/6
(2), 7/7 (1), and 5/7(1). Vertical dorsals
range between 17 and 26 (mean 21.1), hor-
izontal dorsals between 19 and 30 (24.6),
and ventrals between 19 and 32 (24.7). Of
28 adult males, six have moderate nuchal
crest scales and 22 have these scales low;
of 11 females, all have the nuchal crest
scales low. Body crest scales are moderate
in two males and low in 26, and 11 females
have the body crest scales low. The sub-
oculars are separated from the supralabials
in 45 of 49 instances, and contact between
these scales occurs in four lizards (8 per-
cent ) .
The southern specimens from the Port-
au-Prince region and including two from
HisPANioLAN Giant Anole • Schwart::.
103
Mirebalais, consist of ten adult males, three
adult females, and two juvenile females
(MCZ 60013-14). The latter two speci-
mens (with snout-vent lengths of 89 and
92 mm) can be easily dismissed in that
they are presently patternless green. Color
notes in life on one southern male (ASFS
V9024) state that in the green phase, the
lizard had the dorsum a mixture of pale
green, brown, and yellow, with green the
predominant color, the net effect being one
of bead work. The upper surface of the
head was a mixture of pinkish and yellow
scales, the mental region and adjacent up-
per labials were dull yellow, and the venter
pale green. The dewlap was peach with
the dewlap scales yellow. All males (with
the exception of MCZ 69404, which is an
albino ) have some black to dark brown oc-
cipital blotching, usually extending onto
the neck and thence onto the area above
the forelimb insertion. The extent of the
dark anterior markings is variable, but they
are present in all males and quite vivid in
freshly taken specimens. A pale subocular
crescent is present, often extending posteri-
orly to form a pale preauricular blotch,
bounded above by a large dark temporal
blotch that may form an occipital chevron
by joining its mate on the other side.
Southern females were recorded in life as
pale to bright green without any dark pat-
tern, with a moderately well-defined pale
subocular crescent that may expand into a
pale preauricular blotch; the venters were
yellow-green. In one female, the dewlap
was reported as blue-gray with yellow-
green streaks. The hindlimbs are not prom-
inently banded. Neither sex has the throat
marked with any blotching or dotting, al-
though the females have the throat some-
what clouded with dull dark green.
The series for the Sierra de Neiba and
the Cordillera Central consists of six males,
four females, and eight juveniles and sub-
adults. The males were described in life
as being dark green with pale green cross-
bands, or pale green with four paler green
crossbands, or patternless green. The up-
per surface of the head was creamy tan.
black blotches occur in this series of males
also, but the occiput lacks clearly defined
black areas, and the black on the body is
much more extensive than it is in all south-
ern males, the extreme condition being that
shown by ASFS V31395, which has exten-
sive black blotching over two-thirds of the
back and sides. The pale subocular cres-
cent is very obscure, but there is a promi-
nent pale preauricular spot in most males.
The females from this region are plain
green, without dark markings, and there is
a prominent pale supralabial blotch in the
area which in males is occupied by the pale
preauricular blotch. Dewlaps in males and
females were invariably recorded as peach,
and both sexes had charcoal eyeskins. As
in southern specimens, the chins and
throats are pale green and without any defi-
nite markings, except that the throats of fe-
males are sufi^used with darker green. The
eight juveniles and subadults range in
snout-vent length from 68 mm to 92 mm.
The two smallest specimens, a male and a
female, were rich pea-green in life with
four, narrow, cream transverse crossbars,
and the smaller had in addition black
streaking in the green areas and a black
postauricular smudge. The ventral color
was rich pea-green and the dewlap skin
was blue-black. All juveniles and subadults
with snout-vent lengths of between 76 and
92 were bright emerald-green dorsally and
without any dorsal dark or pale markings;
one female juvenile ( snout-vent length 78 )
had a lateral black nuchal spot followed by
a bright yellow preaxillary bar, as well as
a bright yellow subocular mark. The dew-
lap was recorded as black in a juvenile fe-
male with a snout-vent length of 89. Of
the subadults, the most peculiar is a male
(ASFS V31323) with a snout-vent length
of 90 which shows, as preserved, a vague
series of vertical lateral pale and dark
areas, but as yet no black blotching typical
of adult males.
The northern specimens are six males and
one female from Haiti, five males and two
females from the Republica Dominicana,
and one Haitian subadult and two Domin-
104 Bulletin Museum of Comparative Zoology, \o\. 146, No. 2
ican subadult and juvenile lizards. Haitian
males are not only quite different from
northern Dominican males, but they are
also strikingly different from central and
southern males. In the Haitian males, the
dorsum is gray-green with yellow-green
flecking, or a beadwork mixture of dark and
light green scales. The upper surface of the
head is dark with light flecking, and in one
male the head was recorded as dark brown
with the centers of the scales pale purple.
No male has any occipital dark blotching,
and any body blotching, if it is at all pres-
ent, is extremely restricted and maximally
expressed as small black areas above the
forelimb insertion ( MCZ 66147). The pale
subocular crescent is obscure, but there is
a pale postlabial line leading to the auricu-
lar opening. Northern Dominican males,
on the other hand, are brightly colored and
have extensive black neck and side mark-
ings; 'n two males these latter extend far
posteriorly on the body and tend to delimit
two lateral stripes on each side. The upper
surfaces of the head are not mottled but
are pale uniform tan. In life, the pale sub-
ocular crescent is bold and pale blue to
white, and it may extend to the auricular
opening. In Haitian males, the dewlap is
grayish to yellowish peach (pi. 12C5; all
color designations from Maerz and Paul,
1950), pale gray-green (about pi. 19B2),
or yellowish gray (about pi. 20B1). In
northern Dominican males, the dewlap is
pale peach to pale yellow or grayish yellow,
and the dewlap may be speckled with
brown basally.
The single Haitian female is presently
unmarked green, with faint scattered cream
flecking. The larger of the two Dominican
females was pale green above with a darker
brown reticulum outlining a pair of green
lateral stripes on each side. There was a
postauricular brown smudge, followed by a
pale blue axillary smudge. The temples
were yellow-green, the lores pale blue and
brown, the eyeskin pale green, and there
was a pale blue subocular crescent that ex-
tended into a preauricular pale blotch. The
top of the head was marbled pale tan and
dark brown, and the venter was the same
color as the dorsum. The other Dominican
female was green without any dorsal mark-
ings.
The Haitian subadult (MCZ 66148) is a
female with a snout-vent length of 106. It
is speckled with pale scales on a dark
ground like Haitian males. The smaller of
the two Dominican males (ASFS V18008)
has a snout-\'ent length of 75 and was
bright yellow-green above with two cream
crossbands and a yellow subocular cres-
cent. The second Dominican male (ASFS
V32160) has a snout-vent length of 103
mm, and, like Dominican adult males, has
extensive black blotching on the head,
neck, and almost the entire dorsum. The
ground color was pale green, and the dew-
lap was dark brown.
To summarize all the above data, it is
obvious that I have included several popu-
lations in A. r. ricordi which differ rather
strikingly among themselves. Southern
Haitian males are marked with black on
the occiput, neck, and anterior sides, and
central Dominican and northern Domini-
can specimens increase this tendency to
show even more extensive black lateral
markings. On the other hand, northern
Haitian males as a group show very little
or no black markings and are basically
green-flecked green lizards. Northern Do-
minican males are much more colorful than
specimens from elsewhere, and much more
contrastingly marked. On the other hand,
all females are fairly similar, with the ex-
ception of the remarkably colored and pat-
terned female from the northern Republica
Dominicana. I suspect that it will ulti-
mately be shown that there are at least two
more nameworthy populations included in
A. r. ricordi as here defined by me: a north-
ern Haitian subspecies, a northern and cen-
tral Dominican population, as well as the
southern Haitian one. But the specimens
at this time are from such disjunct localities
and are so limited in number that I am un-
willing to make the suggested nomencla-
tural additions.
Remarks. A. r. ricordi occurs in a wide
HisPANioLAN Giant Anole • Schwartz
105
variety of situations but is of course always
associated with trees. Its altitudinal range
is from sea level' at many localities to eleva-
tions of 3500 feet (1068 meters) in the
Chaine de Marmelade in northern Haiti
and 3400 feet (1037 meters) in the Sierra
de Neiba. Almost all specimens taken by
myself and parties were secured sleeping
at night. \\^illiams (1965: 2-3) noted that
in the Monte Cristi region these lizards
sleep in viny tangles, especially where there
are dense "mats" or "curtains" of vines un-
der a canopy. Such a situation is ideal in
the xeric forests in the Monte Cristi area.
At Las Matas de Farfan, the lizards were
easily secured at night in a high-canopied
cafetal, sleeping on limbs, branches, or on
vines, either vertical or horizontal. A speci-
men from Morne Calvaire near Petionville
was seen during the late morning on a
mango tree in an open pasture, about 4 feet
(1.2 meters) above the base. Thomas com-
mented in his field notes upon a specimen
from Le Borgne which was observed 8 feet
( 2.4 meters ) above the ground on the trunk
of a tree; this male led the pursuers a merry
chase through a series of three trees and
finally sought refuge in dense grass on the
ground, where it was caught! The male
from Terrier Rouge was collected with a
slingshot while it rested head-down on the
main branch of a large tree 15 feet (4.6
meters) above the ground. South of Las
Matas de Farfan I secured a juvenile sleep-
ing on a horizontal vine in a tree-fern
thicket adjacent to a mountain brook. The
association of A. r. ricordi with rivers or
lakes is certainly fortuitous; the greatest
concentrations of these lizards occur in such
obviously mesic situations only because
there is often gallery forest restricted, or
limited by man, to streamsides. However,
such a situation is not a guarantee of secur-
ing specimens. In our effort to narrow the
previously existing gap between ricordi and
haleatus in the northwestern Republica
Dominicana, we questioned natives con-
cerning the occurrence of saltacocotes in
the region along the gallery-forested Rio
Yaque del Norte, which here passes
through cactus desert. W'e were assured
that the lizards indeed occurred there, and
we were fortimate in finding a superb area
of gallery forest in the steep-sided valley of
the Rio Guarabo, west of Los Qucmados.
The Guarabo is a southern affluent of the
Yaque, and we had no doubt that these
splendid hardwoods harbored A. ricordi.
But it was not until our fifth nocturnal visit
that a single subadult was secured, despite
the attentions of four collectors. The woods
here present a perfect aspect for A. ricordi
— dense and large trees connected and in-
terlaced with vines and lianas, all quite
rich and mesic; still, our experience indi-
cates that, at least at the time of our visit,
A. ricordi was distinctly uncommon or diffi-
cult to see in what elsewhere surely would
have been a typically simple area for col-
lection of these lizards. In this instance,
demands for at least one specimen from this
region forced persistence which ultimately
yielded the requisite animal. Such may
well be the case in many otherwise xeric
regions, where A. ricordi is restricted to
(and perhaps is rare in) more mesic river-
ine hardwood stands.
Specimens examined. HAITI: Dcpt. de
rOuest, Source Leclerc, Morne Decayette
(MCZ 65729-31); Diquini (MCZ 8619,
USNM 118902, USNM 123347, USNM
123988); Port-au-Prince (AMNH 49501);
Petionville (MCZ 60013-14); Morne Cal-
vaire, 1 mi. (1.6 km) SW Petionville, 2300
feet (702 meters) (ASFS X1711, ASFS
V8514, ASFS V9024); Mirebalais (MCZ
69404); Lancironelle, nr. Mirebalais (not
mapped) (MCZ 68479); Dcpt. de I'Arti-
honite, 8-9 km W Marmelade, 3500 feet
(1068 meters) (ASFS V9925); Dcpt. du
Nord Quest, Port-de-Paix (MCZ 63338);
Dcpt. du Nord, 3 mi. (4.8 km) SW Le
Borgne (ASFS V10005); 2 mi. (3.2 km)
SW Cap-Haitien (ASFS \'10766); Ti
Guinin, nr. Cap-Haitien (not mapped)
(MCZ 66147-49); 8 mi. (12.8 km) E Ter-
rier Rouge (ASFS V10169). RFPOBLICA
DOMINiCANA: Monte Cristi Province, 1
km W Copey (ASFS V1269, ASFS V1411-
12, ASFS V1470); Laguna de Salodillo, 7
106 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
km E Pepillo Salcedo (ASFS V1413);
Dajahon Province, Restauracion (ASFS
V18006-08); Santiago Rodriguez Province,
1.8 mi. (2.9 km) W Los Quemados, 500
feet (153 meters) (ASFS V32160); La Es-
trelleta Province, 6.7 mi. ( 10.7 km ) E
Hondo Valle, 2500 feet (763 meters)
(ASFS V31428); 11.0 mi. (17.6 km) S
Elias Pina, 3400 feet (1037 meters) (ASFS
V31509); San Juan Province, 4.9 mi. (7.8
km) NW Vallejuelo, 2400 feet (732 meters)
(ASFS V31305); 6.1 mi. (9.8 km) S Las
Matas de Farfan, 1800 feet (549 meters)
(ASFS V14562, ASFS V31469, ASFS
V31319-26); 4.1 mi. (6.6 km) NW Juan de
Herrera, 1600 feet (488 meters) (ASFS
V3139.5-99).
Anolis ricordi leberi Williams
AnoUs ricordii leberi Williams, 1965. Breviora,
Mus. Comp. Zool., No. 232: 4.
Tijpe locality. Camp Perrin, Departe-
ment du Slid, Haiti; holotype, MCZ 80935.
Definition. A subspecies of A. ricordi
characterized by the combination of mod-
ally 4 snout scales between second canthals,
6 vertical rows of loreal scales, 3 scales be-
tween the supraorbital semicircles, 4/4
scales between the interparietal and the
supraorbital semicircles, low number of
vertical dorsal scales (14-21; mean 16.5),
low number of ventral scales ( 15-28; mean
20.2), nuchal crest scales usually moderate,
rarely low in males, usually low, occasion-
ally moderate in females, subocular scales
in contact with supralabials in almost 50
percent of the specimens; males either pale
yellow-green with four dark saddles and a
bluish green flank stripe, or with about
three longitudinal dark brown lateral
stripes, or simply dark brown, females
bright green (much brighter than males),
with longitudinal black lines indicated and
at times a greenish tan middorsal wash;
dewlap bright orange or orange with an
anterior brown wash in males, and dull or-
ange, at times suffused or marbled with
brown, in females.
Distribution. Known only from the vi-
cinity of the type locality and Marceline,
on the southern slopes of the Massif de la
Hotte, between elevations of 1000 and 1220
feet (305 and 372 meters), Dept. du Sud,
Haiti.
Discussion. In contrast to the situation
in A. r. ricordi, A. r. leberi is known from
a long series of specimens all from the same
general area, at elevations between 1000
feet and 1220 feet (305 and 372 meters).
Williams (1965: 6) assigned a single juve-
nile (MCZ 38277) from Tardieu, near Pic
Macaya, Dept. du Sud, Haiti, to leberi
with some reservation. This locality is
northwest of Camp Perrin-Marceline, is on
the northern slopes of the Massif de la
Hotte, and is much closer to the known
distribution of the next subspecies to be
described below.
The series of 54 A. r. leberi shows the
following variation. The largest male
(ASFS X3034) has a snout-vent length of
147, the largest female (AMNH 98723)
153; both are from Camp Perrin. Snout
scales at level of the second canthal are ex-
tremely variable, and range between 2 and
7; the mode is 4 (23 specimens). The ver-
tical loreal rows vary between 5 and 8, with
a mode of 6 (26 specimens ) . There are be-
tween 1 and 4 scales between the supraor-
bital semicircles ( mode 3 ) . There are mod-
ally 4 scales between the intei-parietal and
the semicircles; 4 scales are involved in 64
percent of the combinations; actual counts
are 3/3 (3), 3/4 (7), 4/4 (25), 4/5 (10),
5/5 (2), 5/6 (2), 6/6 (1), 3/5 (1), and 5/7
( 1 ) . Vertical dorsals range between 14 and
21 (mean 16.5), horizontal dorsals between
15 and 24 (18.0), and ventrals between 15
and 28 (20.2). Of 39 adult males, 30 have
the nuchal crest scales moderate and nine
have them low; of 13 females, five have the
nuchal scales moderate and eight have them
low. Body crest scales are moderate in 12
males and low in 27 males, whereas only
one female has the dorsal crest scales mod-
erate and 12 have them low. The subocu-
lars are separated from the supralabials by
1 row of scales in 28 specimens and are in
contact with the supralabials in 26 speci-
mens. A. r. leberi is the only population
I
HisPANioLAN Giant Angle • Sclnvnrfz 1U7
that has sucli a high incidence (48 percent) from tlie type locahty. The smallest (MCZ
of subocular-supralabial contact. 83982 ) is a female with a snout-vent length
Males show three basic patterns: 1) dor- of 52. The body is longitudinally streaked,
sal ground color pale yellow-green with but there are as yet no definite longitudinal
four dark brown saddles and a bluish green lines. The subadults ( MCZ 80949-50, a
flank stripe that is complete; 2) about three male with a snout-vent length of 105, and
longitudinal dark brown stripes, the ecu- a female with a snout-vent length of 93)
tral one being the most prominent and both show indications of the longitudinal
complete; 3) and a uniform dark velvety stripes that are characteristic of adults, but
brown. In the two lighter phases, the eye- the stripes are better defined in the sub-
skin is pale blue, chin and throats are dull adult male than in the female. The two
yellow-green, and the subocular crescent is adult males and two adult females from
pale blue and very conspicuous. The dew- Marceline agree in all pattern details with
laps in males are orange (brighter than any the topotypical series; Marceline and Camp
Macrz and Paul designation) or orange Perrin are separated by about 4.5 kilome-
with an anterior brown wash. Females are ters airline.
bright green dorsally (much brighter than Comparisons. Although both A. r. ri-
males) with longitudinal black lines indi- cordi and A. r. leheri have several features
cated. There is a greenish tan wash on the in common, namely, the moderate to low
upper surface of the head, and there may nuchal and body crests, the presence of
be a greenish tan middorsal zone on the some sort of black body markings, and a
body. The dewlap in females is dull or- prominent pale subocular crescent, these
ange, often suffused or marbled with two subspecies are eminently distinct,
brown, and the eyeskin is green, paler than They differ in: modal number of second
that of the dorsum. In males, the venter is canthal snout scales (ricordi 7, leheri 4),
pale green and may be washed with brown modal number of loreal rows ( ricordi 7,
even in the green phase, and in females leheri 6), modal number of scales between
the venter is pale yellow-green, paler than the interparietal and supraocular semicir-
the bright green of the dorsum. cles {ricordi 5/5, leheri 4/4), higher means
In general aspect, male A. r. leheri are of vertical dorsal scales and ventrals (21.1,
lineate dorsally and laterally, the bold dark 24.7 in ricordi, 16.5, 20.2 in leheri, respec-
longitudinal lines usually interrupted by tively) and the very high incidence of con-
four irregular pale vertical crossbands, tact between the subocular scales and the
which are in tinn bordered with darker supralabials in leheri versus the rarity of
pigment. Although my field notes indicate this condition in ricordi. In addition, the
that there are about three longitudinal dark dewlap in male ricordi is most often some
stripes in males, these three stripes are the shade of peach (although the variation in
result of modification of two stripes, of dewlap shades and colors in ricordi is read-
which the more dorsal is the broader. In ily acknowledged), whereas in male leheri
many specimens, this upper flank stripe the dewlap is orange or orange with a
maintains its integrity, but in many others brown anterior wash. A ready hallmark be-
the upper stripe is hollowed centrally, re- tween the two subspecies is the presence of
suiting in three narrow dark stripes, rather a pale preauricular blotch in ricordi, a con-
than two stripes, of which the upper is very dition always absent in both sexes of leheri,
broad and the lower is narrow. Although with the result that instead of the pale sub-
females show some longitudinal striping, it ocular crescent's being incoi-poratcd into a
is much less conspicuous than in males, postlabial line or preauricular blotch as it
Male throats are immaculate, whereas fe- often is in ricordi, it is a bold and contrast-
male throats are suffused with dark green. ing pattern element.
There are three juveniles and subadults Remarks. All Camp Perrin specimens of
108 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
A. r. leheri were collected by natives and
thus I have no precise knowledge of the
habitat nor habits of this subspecies. Camp
Perrin lies in the lower southern foothills of
the high Massif de la Hotte, at about 1000
feet (305 meters), and the area in general
is very mesic and presumably was once
well forested, although now it supports
cafeieres with a high-canopy hardwood
shade cover. Williams (1962: 10) cited
field notes by A. S. Rand and J. D. Lazell,
Jr., on A. r. leheri at Camp Perrin and Mar-
celine; both accounts involve trees associ-
ated with coffee plantings.
Specimens examined. HAITI: Dept. du
Slid, Camp Perrin (ASFS X3033-,35, ASFS
X3038-39, ASFS X3041-42, ASFS X3182,
AMNH 93713-36, MCZ 80935-37, MCZ
S0939-42, MCZ 80944-53, MCZ 83982);
Marceline (MCZ 121115); Marceline area,
ca. 1000 feet (305 meters) (MCZ 122269,
MCZ 121779-80).
Anolis ricordi viculus new subspecies
Holotype. USNM 193974,^ an adult
male, from Castillon, 2500 feet (763 me-
ters), Departement du Sud, Haiti, taken by
native collector on 2.5-26 June 1971. Orig-
inal number ASFS V25059.
Parotypes. ASFS V25058, same data as
holotype; ASFS V25060, same locality and
collector as holotype, 27 June 1971; ASFS
V24801, ca. 2 km (airline) S Castillon,
3500-4000 feet (1068-1220 meters), Dept.
du Sud, Haiti, R. Thomas, 24 June 1971;
ASFS V9335, ca. 5 km (airline) SE Marche
Leon, 2200 feet (671 meters), Dept. du
Sud, Haiti, native collector, 15 March 1966;
MCZ 119035, Castillon, Dept. du Sud,
Haiti, T. P. Webster, A. R. Kiester, and na-
tive collectors, 31 August 1969.
Definition. A subspecies of A. ricordi
characterized by the combination of mod-
ally 6 snout scales between the second can-
thals, 7 vertical rows of loreal scales, 4
scales between the supraorbital semicircles,
4/4 scales between the interparietal and
the supraorbital semicircles, low number of
vertical dorsal scales (15-19; mean 16.7),
moderate number of ventral scales ( 19-24;
mean 21.8), nuchal crest scales usually
moderate but occasionally low in both
sexes, dorsal body crest scales low in both
sexes, subocular scales almost always sep-
arated by one row of scales from suprala-
bial scales; males bright green dorsally
with powdery pale blue-green lateral
stripes, throat pale green and unmarked,
venter pale green with pinkish and yellow-
ish suffusions, females dark olive-green to
bright green with two purple to powdery
blue-gray flank stripes edged with dark
brown, lower sides spotted bright green,
yellow-green, or bright green with four
bright yellow-green crossbands edged with
black, throat pale green; dewlap deep yel-
low to orange in males, dull orange (al-
most brown) to deep yellow with orange
streaking and bluish edge in females.
Distribution. Known only from the vi-
cinity of Castillon on the northern slopes
of the Massif de la Hotte at elevations be-
tween 2200 and 4000 feet (671 and 1220
meters) on the Tiburon Peninsula in Haiti;
probably the subspecies occurring at Tar-
dieu near Pic Macaya (see discussion).
Description of holotype. An adult male
with a snout-vent length of 143 and a tail
length of 165 (regenerated); snout scales
at level of second canthals 6, 7 vertical
rows of loreal scales, 3 scales between su-
praorbital semicircles, 6/5 scales between
interparietal and supraorbital semicircles,
vertical dorsals 15, horizontal dorsals 22,
ventrals 20, one row of scales between sub-
oculars and supralabials, fourth toe lamel-
lae on phalanges II and III 31, nuchal crest
scales moderate, body crest scales low; in
life, bright green above with a pair of lat-
eral stripes on each flank powdery pale
blue-green, the same color also on the face;
throat and neck pale blue-green; venter
pale green with pinkish and yellowish suf-
fusions; dewlap deep yellow, almost or-
ange.
Variation. The series of three males and
three females shows the following varia-
tion. The largest male (ASFS V25058) has
a snout-vent length of 148, the largest
female (ASFS V25060) 141; both are from
HisPANioLAN Giant Angle • Schwartz
109
Castillon. Snout scales at level of the sec-
ond canthal range between 5 and 9; the
mode is 6 (four, specimens). The vertical
loreal rows are 6 or 7, with a mode of 7
( five specimens ) . There are be^^veen 3 and
5 scales between the supraorbital semicir-
cles (mode 4). There are modally 4 scales
between the interparietal and the supraor-
bital semicircles; 4 scales are involved in 58
percent of the combinations; actual counts
arc 4/4 (3), 4/5 (l),5/5 (1), and 5/6 (1).
X'ertical dorsals range between 15 and 19
(mean 16.7), horizontal dorsals between 17
and 27 (20.0), and ventrals between 19
and 24 (21.8). Of three males, two have
the nuchal crest scales moderate and one
has them low; the same situation applies
to the three females. All specimens have
the body crest scales low. The suboculars
are usually separated from the supralabials
by one row of scales and are in contact
with the supralabials in one individual ( 17
percent ) .
Thomas's field notes on three males show
the variation in dorsal coloration and pat-
tern. The dorsum was bright green with
the flank stripes powdery pale blue- green,
this color occurring also on the face. The
throat and neck were also pale blue-green
and the venter was pale greenish with pink
and yellow suffusions. One male (ASFS
V9335) also had a white shoulder patch,
but other pattern details on this individual
were lacking since the specimen was badly
damaged. The dewlap in the males was re-
corded as deep yellow ( almost orange ) and
orange (PI. 11L6). One female was green
to dark olive-green dorsally with two pur-
ple flank stripes, edged with dark brown,
which were powdery blue-gray anteriorly.
The lower sides were spotted and suffused
with bright green or yellow-green. The
venter was pale green with a pinkish wash
in the pectoral region. The second female
was marked in quite a different fashion,
and the specimen still maintains the pattern
after preservation. The dorsum was bright
green with four bright yellow-green trans-
verse body bands with black edges; in this
specimen longitudinal stripes were also
present but only in the nuchal region, and
the venter, including the throat, was pale
green. In both females, the dewlaps were
recorded as "very dull orange" and "deep
yellow, almost brown, anteriorly, with or-
angish longitudinal striae, each edged with
dark gray-green, between striae pale gray-
green and most basal striae greenish; edge
of dewlap grayish (faintly blue)."
Comparisons. A. r. viculus is so very
different from A. r. ricordi in both color
and pattern that detailed comparisons are
hardly necessary. The black occipital, nu-
chal, and anterior body blotches of male
ricordi are absent in male viculus, and the
longitudinally striped pattern in both sexes
of viculus does not occur in ricordi. The
two subspecies differ also in scale counts,
as follows: modal number of snout scales
at second canthals ( ricordi 7, viculus 6 ) ,
scales between interparietal and supraor-
bital semicircles {ricordi 5/5, viculus 4/4),
and much lower means of vertical dorsal
and ventral scales (21.1, 24.7 in ricordi,
16.7, 21.8 in viculus, respectively). The
two taxa are similar in number of loreal
rows, number of scales between the semi-
circles, and in relative frequency of contact
between the subocular and supralabial
scales.
In every way, viculus is much closer to
leheri than to ricordi. The basic pattern
elements are comparable in these two sub-
species, since both are lineate; however,
the longitudinal flank stripes in leheri are
dark, whereas in viculus they are light; the
single banded female viculus is quite dif-
ferent in general aspect from banded le-
heri. As far as scale counts are concerned,
the two subspecies differ in the following
manner: modal number of snout scales at
second canthals {leheri 4, viculus 6), num-
ber of vertical loreal rows {leheri 6, vicu-
lus 7), and scales between supraorbital
semicircles {leheri 3, viculus 4). In mean
number of vertical dorsals and ventrals, the
two subspecies are very similar, and both
have the 4/4 condition as the mode for the
interparietal-semicircle relationship.
Discussion. Williams (1962: 7-8) con-
110 BuUetm Museum of Comparative Zoology, Vol. 146, No. 2
sidered the four specimens then available
from the central portion of the Tiburon
Peninsula as intergrades between ricordi
and leheri. A few more specimens have ac-
cumulated since that time; now there are
one adult male, one juvenile male, and six
adult females from this central region, as
follows: HAITI: Dept. du Sud, Pemel, nr.
Miragoane (not mapped) ( MCZ 66015-
16), PaiUant, 1800 feet (549 meters)
(ASFS V26535-37); Fond des Negres
(ASFS V26254, USNM 72631, USNM
72633). As preserved, the adult male
shows fairly obvious longitudinal streaking
of gray and dull green, a few scattered
dark flecks or small blotches above the
forelimb insertion, and a prominent suboc-
ular pale crescent. The adult male is an
almost ideal representation of extreme in-
tergradation between viculus and ricordi,
with both pale longitudinal stripes and
scattered remnants of the typical ricordi
extensive body blotching. Three recently
collected females in life were green with
longitudinal stripes, which were delimited
by absence of black flecking that occurs
elsewhere on the green ground. In the
brown phase, these longitudinal stripes had
a reddish wash. In all females, the pale
subocular crescent is obvious and bold, and
in one female (MCZ 66016) there is an ad-
ditional preauricular pale area that resem-
bles the condition in nominate ricordi. I
have no color data on the male dewlaps,
but that of one female (ASFS V26535) was
dull yellow distally and pale blue, smudged
with charcoal, basally; the dewlap scales
were yellow-green. The juvenile male
(snout-vent length 79) in life had a pat-
tern of longitudinal dorsolateral stripes and
dorsal crossbands, with a pale yellow sub-
ocular crescent. I interpret these lizards as
intergradient between ricordi and viculus.
The central Tiburon localities, however,
are far removed from the known localities
of viculus (110 km) on one hand and of
ricordi (70 km) on the other. Williams
(1965: 7) regarded the Fond des Negres
and Pemel specimens as ricordi X leheri
intergrades, and they could indeed be so
interpreted. Since, however, leheri occurs
on the southern slopes of the Massif de la
Hotte, and viculus on the northern slopes
of that range, and since all intergradient
specimens are from the northeastern re-
gions of the extreme eastern portion of the
Massif de la Hotte, it seems much more
likely that these central Tiburon specimens
are intergradient between ricordi and vic-
ulus on geographic grounds. They do not
disagree with my concepts of how inter-
grades between these two subspecies prob-
ably should appear.^
1 Since the above comments on the intergradi-
ent specimens were written, Williams has secured
a series of 28 lizards (MCZ 132302-29) from St.
Croix, 1 mi. (1.6 km) from Paillant, Dept. du
Sud, Haiti, from this same general region. There
are no color data on the specimens. The measure-
ments ( in mm ) and scale counts of these lizards,
combined with those from the eight previously
available soecimens, follow. Largest male ( MCZ
132325) 155, largest female (ASFS V26535) 148.
Snout scales at second canthals 4-9 (mode 6);
loreal rows 4-8 ( mode 6 ) . Modally 4/4 scales
between the interparietal and the supraorbital
semicircles; other counts: 3/3 (2), 3/4 (2), 4/5
(4), 5/5 (11), 5/6 (1), 3/5 (1); 4 scales are in-
volved with 50 percent of the combinations. Ver-
tical dorsals range between 14 and 21 (mean
17.0), horizontal dorsals between 15 and 24
(18.3), and ventrals between 16 and 31 (21.4).
Of the males, four have the nuchal crest scales
moderate and 15 have them low; of the females,
one has the nuchal crest scales moderate and 15
have them low. Body scales are low in all adult
specimens. The suboculars are modally separated
from the supralabials by one row of scales and
are in contact with the supralabials in seven liz-
ards ( 19 percent).
In scale counts, the entire series is much closer
to viculus than to ricordi; however, in some char-
acteristics, the series is closer to lebcri or to the
subspecies yet to be described from extreme south-
eastern Haiti. In fact, comparison of the scale
counts shows that there is little resemblance be-
tween the modes and means between these geo-
graphically intermediate specimens and nominate
ricordi, and as a whole they seem much more
closely allied to one of the other Tiburon subspe-
cies.
The males in the St. Croix series are variable
in pattern but none shows any clear-cut dark
blotching, typical of A. r. ricordi. Some males
are more or less unicolor (medium brown as pre-
served), whereas others have longitudinal stripes,
alternating light and dark, with usually one broad
HisPANioLAx Giant Angle • Schtvartz
111
There remains one other specimen from
the distal portion of the Tibnron Peninsnhi;
this is a jnvenile male ( MCZ 38277) with
a snont-vent length of 7(S, from Tardieu,
near Pic Macaya, collected by P. J. Darling-
ton. It is presently dull brown, but there
are clear indications of black-edged dorsal
crossbands that closely resemble the condi-
tion in one of the female paratypes of vi-
culus. Tardieu is presently unlocatable on
modern maps, but Darlington has indicated
to Williams that this place lies just to the
north of Pic Macaya, and thus rather close
to Castillon. Since there are no juvenile
dark stripe along the upper sides and most prom-
inent. One male has extensive dark brown body
markings, vertically oriented and alternating with
paler tannish areas to give a more-or-less verti-
cally barred appearance. The pale subociilar
crescent is very obvious in all males, and there is
no indication of a pale preauricular area.
The females are undistinguished. Most are
more or less solid green with some scattered paler
green scales to give a beadwork effect dorsally,
but there are also indications in some specimens
of longitudinal paler areas to give a somewhat
longitudinally lined appearance. As in the males,
the subocular pale crescent is obvious, and there
may be a weakly differentiated pale preauricular
area.
This newly collected series of A. ricordi is puz-
zling. The entire lot is so like, in general aspect,
specimens of leberi and viculus (and the yet-to-
be-named subspecies in southwestern Haiti ) and
shows so little tendency toward A. r. ricordi that
it is difficult to interpret them as intergradient be-
tween ricordi and viculus. The adult male (MCZ
66015) noted above in the body of the text is
from "Pemel, near Miragoane," a site that is un-
locatable on modern maps. Pemel may be "near
Miragoane" in only the very broadest sense. Spec-
imens that are known to have been taken in the
Miragoane-Paillant area show little evidence of
intergradation between vicidus and ricordi, and
are much closer to the former subspecies.
Everything considered, I strongly suspect that
with additional collecting on the central Tiburon
Peninsula it is probable that two taxa will be
found to occur here in sympatry and without
wide areas of intergradation, or that ricordi-styled
anoles occur close to ( but do not intergrade with )
leberi-styled anoles. The evidence at the moment
is far from unequivocal that ricordi and viculus
intergrade in this area. Only much additional
collecting along the Tiburon Peninsula will reveal
the actuality of the relationships of A. ricordi with
its southeastern relatixes.
viculus, I have no concepts of their appear-
ance; on geographic grounds, however, I
have little doubt that the Tardieu male is
representatixe of viculus rather than of
leberi.
Remarks. It may seem remarkable that
there should be two distinctive subspecies
of A. ricordi in such close geographical
proximity; Castillon and Marceline are sep-
arated by only 29 kilometers airline, and if
the Tardieu specimen is viculus, then the
distance between the localities for the two
subspecies is even shorter. However, be-
tween Castillon and Marceline lies the high
ridge of the La Hotte, including the cul-
minating peak of that range, Pic Macaya,
with an elevation of 7698 feet (2347 me-
ters). Such high and rugged country is
probably ecologically unsuitable for A. ri-
cordi, and the northern and southern pop-
ulations associated with the La Hotte have
differentiated because of isolation caused
by the intervening massif.
The known altitudinal range of A. r. vic-
ulus is between 2200 and 4000 feet (671
and 1220 meters). The Castillon area, ac-
cording to Richard Thomas, is generally
mesic but much of the original forest has
been cut. Still, enough trees and ravine
woods remain to offer haven for such a
tolerant and adaptable species as A. ricordi.
One female from south of Castillon was
taken by Thomas on the trunk of a large
tree about 5 feet (1.5 meters) above the
ground; all other ASFS specimens were se-
cured by natives.
The name vicidus is from the Latin for
"hamlet" or "small village" in allusion to
Castillon, the type locality.
Anol'is ricordi subsolanus new subspecies
Holotype. MCZ 130270, an adult male,
from Source Carroye, near Saltrou, Depart-
ement de I'Ouest, Haiti, one of a series col-
lected by George Whiteman in March 1972.
Paratypes. MCZ 130264-69, MCZ
130271-77, same data as holotype; MCZ
69405, nr. Saltrou, Dept. de I'Ouest, G.
Whiteman, summer 1962.
Definition. A subspecies of A. ricordi
112 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
characterized by the combination of mod-
ally 5 snout scales between second canthals,
5 vertical rows of loreal scales, 3 scales be-
tween the supraorbital semicircles, 4/4
scales between the interparietal and the
supraorbital semicircles, moderate number
of vertical dorsal scales (16-21; mean
17.3), moderate number of ventral scales
(18-27; mean 21.0), nuchal crest scales
rarely moderate, usually low in males, low
in females, subocular scales always sepa-
rated from supralabials scales by one row
of scales; males vaguely lineate dorsally
with two broad lateral grayish flank stripes,
or with three paler ( green in life? ) cross-
bands; females like males, or heavily
blotched with black laterally and on the
occiput, the black lateral markings in the
areas that are elsewhere occupied by
the gray lateral flank stripes; a pale suboc-
ular crescent present and prominent but no
pale preauricular blotch; dewlap color un-
known.
Distribution. Known only from the re-
gion about Saltrou, in extreme southeastern
Haiti, but see discussion below.
Description of holotype. An adult male
with a snout-vent length of 144 mm and a
tail length of 209 mm ( regenerated ) ; snout
scales at level of second canthals 4, 6 ver-
tical rows of loreal scales, 3 scales between
supraorbital semicircles, 3/4 scales between
the interparietal and the supraorbital semi-
circles, vertical dorsals 18, horizontal dor-
sals 17, ventrals 19, one row of scales be-
tween suboculars and supralabials, fourth
toe lamellae on phalanges II and III 33,
nuchal crest scales low, body crest scales
low; as preserved, dorsum dull dark brcwn
with three prominent blue-green cross-
bands, more or less confluent middorsally,
and outlined in dark brown to black; top
of head brown, paler than sides; throat
greenish, dewlap dull gray; belly dark
gray, underside of hindlimbs green; tail
brown.
Variation. The holotype and paratypic
series are composed of 10 males and five
females. The largest male (MCZ 130274)
has a snout-vent length of 152, the
largest female (MCZ 69405) 150; the male
is a topotype, the female is from near Sal-
trou. Snout scales at level of the second
canthals range between 4 and 7; the mode
is 5 (six specimens). The vertical loreal
rows vary between 5 and 7, with a mode of
5 (eight specimens). There are between 2
and 4 scales between the supraorbital semi-
circles (mode 3). There are modally 4
scales between the intei-parietal and the
supraorbital semicircles; 4 scales are in-
volved in 58 percent of the combination;
actual counts are 3/3 ( 1), 3/4 (3), 4/4 (4),
4/5 (3), 5/5 (2), and 5/6 (2). Vertical
dorsals range between 16 and 21 (mean
17.3), horizontal dorsals between 14 and 23
(17.1), and ventrals between 18 and 27
(21.0). Of 10 males, two have the nuchal
crest scales moderate and eight have these
scales low; all five females have the nuchal
crest scales low. All specimens have the
body crest scales low. In all specimens the
subocular scales are separated from the su-
pralabials by one row of scales.
I have no color notes in life nor have I
seen live specimens of A. r. suhsolanus.
Consequently, my comments on pattern in
this subspecies are based solely upon pre-
served material. In the series of males and
females, each sex shows two basic patterns.
The more common is a pair of longitudinal
flank stripes, the upper being broader, usu-
ally dull grayish in contrast to a greenish
ground color. In two specimens (one male
and one female; MCZ 130267 and MCZ
69405) these stripes are very prominent
and black; although they no longer have
their integrity in the female, they are still
very obvious. In addition, in the female
there is black pigment in the occipital re-
gion. A pale subocular crescent is present
in all specimens and is usually very con-
spicuous. In two specimens (the holotypic
male and a female— MCZ 130266) the dor-
sal pattern consists of three transverse
crossbands that are green, more or less
fused middorsally, and outlined with black
or dark brown. Many females show the lat-
HisPANioLAN Giant Angle • Schtvartz 113
eral flank stripes much less clearly than do
the males, but usually the stripes are at
least indicated. •
Comparisons. In general aspect, sub-
sola nus much more closely resembles far
western leheri and vicuhis than geographi-
cally closer ricordi. The latter subspecies,
however, occurs on the nortliern side of the
Massif de la Selle, whereas the localities for
suhsolamis are to the south of that range.
Since I do not know the coloration in life
of suhsolamis, I am unable to compare its
pigmentation with that of the other subspe-
cies. The presence of both longitudinally
striped and transversely barred specimens
in suhsolamis suggests its affinity with leheri
and vicuhis. A. r. suhsolamis differs from
A. r. ricordi in that the latter has (in its
southern populations) dark anterior mark-
ings on the occiput and above the forelimb
insertions, whereas these markings are ab-
sent in subsolanus. Additionally, southern
ricordi are patternless green, whereas sub-
solaiius females are longitudinally lined and
may have heavy dark anterior markings
(somewhat like male A. r. ricordi). At the
time of Williams's review of A. ricordi
(1965: 2), there was but a single A. ricordi
from the Saltrou region; by chance, this
specimen (MCZ 69405) is the heavily
marked female upon which I commented
above. Although Williams (loc. cit.) con-
sidered it a male, it lacks enlarged postanal
scales and a tail "fin," and it is a female.
Since female A. r. ricordi lack dark anterior
markings, this female is really quite differ-
ent from females of the northern subspe-
cies.
From the western subspecies leheri and
viculus, suhsolamis difi^ers meristically in
the following ways. From leheri, suhso-
lanus differs in having 5 versus 4 snout
scales at the second canthal, 5 versus 6 ver-
tical loreal rows, higher means in vertical
dorsal scales and ventral scales, and also
lacks specimens that have the suboculars in
contact with the supralabials ( leheri has 48
percent of the specimens with this condi-
tion). From viculus, suhsolanus differs in
having 5 versus 6 snout scales at second
canthal, 5 versus 7 vertical loreal rows, 3
versus 4 scales between the supraorbital
semicircles, and higher means of vertical
dorsal scales and ventral scales. In addition
to the pattern differences noted above
which differentiate suhsolanus from nomi-
nate ricordi, suhsolanus has 5 versus 7 snout
scales at the second canthals, 5 versus 7
vertical loreal rows, 3 versus 4 scales be-
tween the supraorbital semicircles, 4/4 ver-
sus 5/5 scales between the interparietal and
the supraorbital semicircles, and lower
means in vertical dorsal scales and ventral
scales.
Remarks. I am once more hampered in
my interpretation of suhsolanus by the
large distributional gap between its two
stations and any other stations for A. ricordi
to the west. The absence of specimens
from the southern coast, from such well-
known areas as Jacmel and Aquin, is truly
puzzling. The nearest locality to suhsolanus
along the Tiburon Peninsula is Fond des
Negres [ricordi X viculus), some 120 kilo-
meters to the west. Still further, the area
known to be occupied by A. r. leheri lies
some 205 kilometers to the west, near the
peninsula's tip. Known stations for A. r. ri-
cordi are very much closer (40 kilometers)
but lie to the north of the Massif de la
Selle. Closer even than' any of these is
harahonae; harahonae and suhsolanus are
known in this region for localities separated
by about 11 kilometers (see comments be-
low), but there is no evidence of intergra-
dation between these two taxa.
It is perhaps pertinent that leheri, vicu-
lus, and suhsolanus all seem closer in most
characteristics to each other than they do
to nominate ricordi. If it were not for the
specimens that I interpret as intergradient
between ricordi and viculus in the Mira-
goane-Paillant-Fond des Negres region, I
would be very tempted to consider these
three taxa as a species distinct from A. ri-
cordi. Much additional material from along
the Tiburon Peninsula will perhaps show
that my interpretation is wrong.
114 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
The name subsolamis is from the Latin
for "eastern," in alhision to the occurrence
of this subspecies in southeastern Haiti.
The precise areas where suhsolanus occurs
are a matter of question. I am unable to
locate Source Carroye on any modern map.
Williams advised me that Source Carroye
is very near Thiotte (according to the col-
lector, "Source Carroye is located northeast
direction and about V2 mile from the main
road after you leave the place of the
'marche'," that market being at Thiotte).
The elevation of Thiotte is about 900 me-
ters. The lone specimen from "near Sal-
trou" also poses the problem of just how
"near" this specimen was taken to Saltrou
itself. Any information on details of local-
ities or elevations of this and other speci-
mens taken along the Dominico-Haitian
border are mandatory. The distance be-
tween the Thiotte locality for suhsolanus
and the Pedernales specimens of harahonae
is about 11 kilometers. It is especially per-
tinent that harahonae is not known, along
the Dominico-Haitian border, from the low-
lands (where, incidentally, Anolis coeles-
tinus is called saltacocote by the natives),
but that harahonae occurs here as an in-
habitant of mesic riverine woods at an ele-
vation of 600 feet ( 183 meters ) .
Anolis barahonoe Williams
Anolis ricordii harahonae Williams, 1962. Brevi-
ora, Mus. Comp. Zool., No. 155: 8.
Ttjpe locality. Polo, Valle de Polo, Bara-
hona Province, Repiiblica Dominicana;
holotype, MCZ 43819.
Defiyiition. A giant species of Hispanio-
lan Anolis characterized by the combina-
tion of moderate size (males to 158 mm,
females to 148 mm snout-vent length),
snout scales at level of second canthal
scales 2 to 5 (mode 4), vertical loreal rows
2 to 5 (mode 6), scales between supraor-
bital semicircles 1 to 4 (mode 2), inteipa-
rietal scale separated from supraorbital
semicircles modally by 4 scales, vertical
dorsal scales generally small ( 15 to 34 in
standard-distance), ventral scales relatively
small (17 to 29 in standard-distance), nu-
chal crest scales in both sexes rarely high,
usually moderate to low, dorsal body crest
scales rarely moderate, usually low, suboc-
ular scales rarely in contact with suprala-
bial scales; dorsal body coloration basically
lichenate gray-green, grays, to browns and
black, giving a blotched effect that also oc-
curs in even the smallest juveniles, and
rarely (only in juveniles) with any indica-
tion of transverse crossbars, or solid brown
to grayish with faintly bluish white dark-
edged ocelli; dewlap pale yellow to peach
in males, pale yellow to pale peach in fe-
males; pale subocular crescent absent in
adults but indicated in juveniles by a pale
subocular spot.
Distrihution. The Sierra de Baoruco
and associated lowlands on the Peninsula
de Barahona, Repiiblica Dominicana, in-
cluding (probably) the semi-xeric forests
of the lowlands south of the Sierra de
Baoruco and southern Haiti; altitudinal dis-
tribution from sea level to 2600 feet (793
meters) northeast of Las Auyamas, Bara-
hona Province.
Anolis harahonae harahonae Williams
Type locality. Polo, Valle de Polo, Bara-
hona Province, Repiiblica Dominicana.
Definition. A subspecies of A. hara-
honae characterized by the combination of
modally 4 snout scales between second can-
thai scales, 4 vertical rows of loreal scales,
2 scales between the supraorbital semicir-
cles, 4/4 scales between the interparietal
and the supraorbital semicircles, relatively
low number of vertical dorsal scales ( 15-
23; mean 17.2), high number of ventral
scales (17-29; mean 22.1), nuchal crest
scales moderate to low, body crest scales
rarely moderate, usually low, subocular
scales usually separated from supralabial
scales by one row of scales, both sexes and
juveniles patterned with varying shades of
gray-green, grays, browns and black, giv-
ing a lichenate blotched effect; juveniles
with vague indications of three transverse
gray bands but that pattern only very
rarely even indicated in adults; dewlap
pale yellow to pale peach in both sexes, the
HisPANiOLAN Giant Angle • Schtvartz 115
female dewlap suffused with gray basally;
pale subocular crescent absent in adults but
indicated by a clear white subocular spot in
juveniles and subadults.
Discussion. A. b. harahonae has a rela-
tively circumscribed range in the Sierra de
Baoruco in the southeastern Republica Do-
minicana. Until our 1971 collections, the
taxon had been known only from the east-
ern portion of that massif, but two speci-
mens taken 13.0 mi. ( 20.8 km ) N of Peder-
nales along the Dominico-Haitian border
are unquestionably A. harahonae. These
individuals differ slightly from more east-
ern specimens of A. h. harahonae in colora-
tion, but they are so close to the nominate
subspecies that for the moment I have no
hesitancy in regarding them as that taxon.
The series of 33 specimens of A. h. hara-
honae shows the following variation. The
largest males (ASFS V29722, MCZ 125504)
have snout-vent lengths of 158, the largest
female (AMNH 50256) 148; the males are
from north of Pedernales and near Polo,
and the female is from Barahona. Snout
scales at level of second canthals vary be-
tween 2 and 5; the mode is 4 (18 speci-
mens). The vertical loreal rows vary be-
tween 5 and 8, with a mode of 6 (11
specimens). There are between 1 and 4
scales between the supraorbital semicir-
cles (mode 2). There are modally 4 scales
between the inteiparietal and the supraor-
bital semicircles; 4 scales are involved with
58 percent of the combinations; actual
counts are 3/3 (3), 3/4 (6), 4/4 ( 13), 4/5
(3), and 5/5 (5). Vertical dorsals range
between 15 and 23 (mean 17.2), horizontal
dorsals between 15 and 24 (18.2), and ven-
trals between 17 and 19 (22.1). Of 16
males, seven have the nuchal crest scales
moderate and nine have these scales low;
of 10 females, three have these scales mod-
erate and seven have them low. Body crest
scales are moderate in one male and low in
15, whereas all 10 females have the body
crest scales low. The subocular scales are
separated from the supralabial scales in 32
of 33 specimens (3 percent).
Exclusive of the male and female from
north of Pedernales, eastern specimens of
A. /;. harahonae are lichenate or blotclied
with gray-green, grays, browns, and black
in a random pattern, although occasional
individuals show remnants of the slightly
more obviously banded condition of the
juveniles. No specimen has been recorded
in the field as being bright green, and in
general the tones of green in harahonae
are dull and grayish. Some specimens (es-
pecially ASFS V30921, a male) were re-
corded as being gray, heavily blotched with
black, and thus without any green tints
whatsoever. The dewlap color in males
varies between pale peach and peach, and
in females between pale peach and yellow.
The Pedernales specimens were recorded
in life as being dark brown to gray dorsally,
obscurely banded with tannish. The heads
were tan above, the eyeskin pale gray, and
the female had the upper surfaces of all
limbs banded green and dark brown. The
most noteworthy difference between these
western specimens and those from the east-
ern uplands of the Sierra de Baoruco and
its associated lowlands is that the dewlaps
in both sexes were pale yellow, that of the
female suffused with gray basally.
Available juveniles and subadults vary in
length between 62 and 95. The juveniles
are colored and patterned essentially like
the adults, except that three pale grayish
crossbands are vaguely indicated in most
specimens. These bands are quite indis-
tinct and much obscured by the lichenate
harahonae pattern. Some juveniles were re-
corded as being banded and mottled pale
gray, dull pea-green, and black, witli a
black nuchal patch and a white nuchal
crescent on each side, whereas others were
recorded as crossbanded gray and dusky,
with some greenish on the lips, and the tails
banded gray and dusky to cream.
Remarks. Specimens secured by myself
and parties have all been taken in wooded
situations, between elevations of 600 feet
and 2600 feet ( 183 and 793 meters). River-
ine woods and the large shade trees in the
upland cafetales of the Sierra de Baoruco
offer optimum habitat for the species. Both
116 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
adults and juveniles were secured sleeping
at night; in general, the juveniles sleep
lower on shrubs and low trees, whereas
adults sleep higher (up to 15 feet — 4.6 me-
ters) on limbs, branches, and woody vines.
At night, despite the absence of bright
greens in the coloration, the lizards are
quite obvious because their pale grayish
hues contrast to the adjacent greenery. All
ages of A. h. harahonae sleep exposed, as
do other Hispaniolan giant anoles. The
pair from 13.0 mi. N Pedernales were se-
cured in rich riverine woods at an eleva-
tion of 600 feet ( 183 meters ) ; this is purely
a gallery forest situation, since in this re-
gion the open slopes are clad in Acacia for-
est or dry scrubby woodlands, whereas
rivers and creeks support much more luxu-
riant arboreal growth.
Almost all localities for A. h. harahonae
are in the highlands. However, the lizard
presumably occurs in coastal forested re-
gions as well. There are specimens from
the city of Barahona ( which is coastal ) and
from halfway between Enriquillo and
Oviedo, which is presumed to be coastal or
nearly so. A third specimen from Enri-
quillo likewise is presumably from a coastal
locality. However, in each of these cases,
it is possible that the lizards were secured
in the adjacent Sierra de Baoruco; this
mountain range comes abruptly to the coast
between Barahona and Enriquillo, and it
would be a simple matter to label speci-
mens from non-coastal localities as having
come from coastal populated areas. Al-
though negative evidence at best, we have
never oiuselves secured A. harahonae along
this coastal region, and residents of Bara-
hona responded negatively when ap-
proached to collect this lizard for us.
A. /;. harahonae is known from a locality
(13.0 mi. N Pedernales) that is only (pre-
sumably) 11 kilometers from a locality
(Thiotte) where A. r. suhsolanus occurs.
There are no other localities where these
two species approach each other, although,
since the northern slopes of the Sierra de
Baoruco are confluent with the northern
slopes of the Massif de la Selle and its affil-
iates, it is not unlikely that somewhere
along these northern reaches A. harahonae
comes into contact with A. r. ricordi. There
is no obvious reason for A. h. harahonae to
be promptly replaced by A. r. suhsolanus
at the Dominico-Haitian border; the polit-
ical boundary on these southern slopes is
the Rio Pedernales, a small stream that
surely offers no obstacle for these arboreal
lizards. It follows that A. h. harahonae
must occur in southeastern Haiti. Thus, as
previously noted, the accuracy of the suh-
solanus localities is more than academic. It
is possible that in southeastern Haiti, A.
harahonae is a more lowland lizard and A.
ricordi (suhsolanus) occurs on the higher
and better forested slopes of the Massif de
la Selle — the division may thus be altitu-
dinal as well as ecological. The precise re-
lationships between these two species re-
main to be determined; only further
detailed collecting in extreme southeastern
Haiti will reveal the siutation there. As far
as distinguishing A. r. suhsolanus from A.
h. harahonae, there is no problem, since the
styles of pattern (and presumably colora-
tion) are so very different as to preclude
confusion. If intergradation between suh-
solanus and harahomie occurs (and since I
here regard harahonae as a species distinct
from ricordi, I am obviously convinced that
it does not), then it must take place very
quickly, in a distance of some 11 kilome-
ters, since suhsolanus and the Pedernales
harahonae are completely different and
typical of their own populations, without
any indication of intergradation between
them.
Specimens examined. REPUBLIC A DO-
MINICAN A: Barahona Province, Barahona
(AMNH 50255-56); 14 km SW Barahona,
1200 feet (366 meters) (ASPS V23460-63,
ASFS V30263-70); Valle de Polo (MCZ
56141, AMNH 51235-37, AMNH 51240,
AMNH 51036); nr. Polo (MCZ 125504-06);
Las Auyamas (ASFS V30921); 8 km NE
Las Auyamas, 2600 feet (793 meters)
(ASFS X9676); Hermann's finca, nr. Par-
aiso (AMNH 51231-33); Enriquillo
(AMNH 51241); Pedernales Province, half-
HisPANiOLAN Giant Angle • Schwartz
117
way between Enriquillo and Oviedo
(AMNH 51230); 13.0 mi. (20.8 km) N
Pedernales, 600 feet ( 1S3 meters ) ( ASFS
V29722-23); locality unkno\\ai (AMNH
51229).
Anolis borahonoe olbocellotus
new subspecies
Holotype. MCZ 125611, an adult male,
from 13.1 mi. (21.0 km) SW Enriquillo,
Pedernales Province, Republiea Domini-
cana, taken by Richard Thomas on 10 De-
cember 1964. Original number ASFS
V4422.
Definition. A sub.species of A. hara-
honae characterized by the combination of
4 snout scales between second canthal
scales, 7 vertical rows of loreal scales, 3
scales between the supraorbital semicir-
cles, 4/4 scales between the interparietal
and the supraorbital semicircles, apparently
relatively high number of vertical dorsal
scales (19), high number of ventral scales
(26), nuchal crest scales high, body crest
scales low, subocular scales separated from
supralabial scales by one row of scales,
male (females unknown) dorsal ground
color nonlichenate brown to grayish with
white (faintly bluish) randomly placed
dark-edged ocelli, head light brown above,
dewlap pale yellow with a pink margin,
and a pale subocular spot.
DistriJnition. Known only from the type
locality, but presumably distributed
through the semi-arid forests of the Penin-
sula de Barahona south of the Sierra de
Baoruco (see discussion).
Description of holotype. An adult male
with a snout-vent length of 150 and a tail
length of 265; snout scales at level of sec-
ond canthal 4, 7 vertical rows of loreal
scales, 3 scales between interparietal and
supraorbital semicircles, vertical dorsals 19,
horizontal dorsals 23, ventrals 23, one row
of scales between suboculars and suprala-
bials, fourth toe lamellae on phalanges II
and III 34, nuchal crest scales high, body
crest scales low; in life, dorsum brown to
grayish, not lichenate, with randomly scat-
tered white (faintly bluish) dark-edged
ocelli involving from 1 to 4 scales; venter
white with gray mottling or stippling; dew-
lap pale yellow with pink along its outer
margin; upper surface of head light brown,
with large pale subcircular areas anterior
to the ear opening, and a conspicuous pale
blotch ])el()w the eye; soles of hands and
feet conspicuously pale yellow.
Comparisons. No mensural nor meris-
tic characters separate aU)ocellatns from
barahoiuie. On the other hand, the distinc-
tive coloration, pattern, and dc^wlap color
of aJhoceUatus are very different from those
of harahonae, and the presence of high nu-
chal crest scales likewise differentiates al-
hocellatus from the moderate to low scales
in Imrahonae. More detailed comparisons
are impossible, but certainly aIJ)OceIl(itus is
quite distinctive when compared with ])ara-
honue.
Discussion. It may seem foolhardy to
name a subspecies of A. harahonae from a
single specimen whose locality is only 11
kilometers from a presumed locality for A.
h. harahonae (half way between Enriquillo
and Oviedo). The holotype of A. h. alho-
cellatus is that lizard about which Williams
(1965: 4) commented, saying it "is typical
in squamation but peculiar in having very
distinct small Ufi,ht spots on the flanks
It will be recalled that it was a specimen
from Enriquillo (AMNH 51241) that
caused some hesitation when harahonae
was first described. In AMNH 51241 the
pattern was thought to be obscure banding;
the present specimen clearly shows spots
tending to be vertically aligned — a condi-
tion which is easily transformed into verti-
cal banding. It is possible that the ricordii
populations in the vicinity of Enriquillo
consistently show a distinctive pattern
though charact(>ristically harahoiuie in
squamation."
The specimen (AMNH 51230) from half-
way between Enriquillo and Oviedo is a
young male with a snout-vent length of
121. Since this lizard presumably came
from the lowlands of the Peninsula de
Barahona, it might logically be expected to
be alhocellatiis. However, the lizard is now
118 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
drab patternless brown, and there are no
indications that it was ever spotted. Pre-
sumably albocellatus and harahonae inter-
grade between Enriqiiillo (which Hes at
the extreme southeastern corner of the Si-
erra de Baoruco) and Oviedo (which Hes
well down on the Peninsula de Barahona).
Several facts have prompted my naming
this lonely specimen. First, I have exam-
ined the Enriquillo specimen noted by Wil-
liams, and, although it shows some indica-
tion of vertical crossbars, they are not any
more conspicuous than those in some more
recently taken A. /;. harahoruie from the
Baoruco highlands (Williams examined
only 17 harahoime at the time of its original
description; I have studied almost twice
this number). Secondly, the xeric to semi-
arid region south of the Sierra de Baoruco
has come to be known as an area of local
differentiation at the subspecific level for a
variety of reptiles; this alone is no reason
for naming albocellatus, of course. Thirdly,
although since 1964 when the holotype was
collected both I and others have spent con-
siderable time on the Peninsula de Bara-
hona and in the vicinity of Oviedo, we have
never seen or secured another A. harahonae
in this region. In September 1966, the very
severe hurricane Inez passed directly across
the Peninsula. What had once been high-
canopied semi-arid forest (as at Oviedo)
has been either totally destroyed or been
reduced (by 1969) to a landscape of bare
snags with some leafy growth just now be-
ginning to appear but at a much lower
canopy-level than previously. The changes
between the Oviedo area in 1964 and 1969
are so massive that, upon my first visit there
after Inez, I was unable to orient myself in
reference to our older collecting localities!
Certainly this entire region has suffered
greatly, and, with the destruction of trees,
it seems reasonable to assume that A. hara-
honae has suffered equally. The population
may never have been high, since such semi-
arid woods are not at all optimal habitat
for any of the Hispaniolan giant anoles, and
the destruction of the habitat must surely
have affected A. h. albocellatus adversely.
Since persistent visits to this area have
yielded no new material, and since the liz-
ard may presently be very rare, I have de-
cided upon the present course rather than
wait in hope for someone to secure a sec-
ond ( or more ) lizard.
Remarks. The Peninsula de Barahona
has been shown to have distinctive subspe-
cies (or even species) of a variety of rep-
tiles. Species that have described endemic
subspecies south of the Sierra de Baoruco
include: Sphaerodacttjhis difficilis Barbour,
Leiocephalus Imrahonensis Schmidt, Am-
eiva chrysolaema Cope, Ameiva lineolata
Dumeril and Bibron, Arnphisbaena gona-
vensis Cans and Alexander, and Dromicus
parvifrons Jan. Endemic Peninsula de
Barahona species are: Anolis longitibialis
Noble, Typhlops sijntherus Thomas, Lep-
totyphlops pyrites Thomas, and Uromacer
ivetmorei Cochran. Only one amphibian,
Eleutherodactylus alcoae Schwartz, is re-
stricted to the Peninsula. To the former
list can now be added Anolis harahonae.
The eastern half of the Peninsula, although
xeric, was originally clothed in dry forest,
much of it upon a series of limestone ter-
races, the highest point of which is the
Loma Gran Sabana, having an elevation of
1082 meters in the north and descending to
Cerro Caballo, and Loma de Chendo, hav-
ing elevations of 322 and 233 meters, re-
spectively, to the south. West of this ridge,
the land descends abruptly to Acacia-cac-
tus desert to the east of Cabo Rojo, and
this habitat continues to the Dominico-
Haitian border at Pedernales. Presumably,
A. b. albocellatus occurs throughout the
eastern half of the Peninsula in the for-
merly high-canopied forests of the lime-
stone terraces.
The holotype was secured by Richard
Thomas during the day in a viny tangle in
semi-xeric woods near Oviedo; the lizard
was in an edge situation, since beyond the
dense vine tangle the woods thinned to
more scrubby and cleared areas.
The name albocellatus is from the Latin
HisPANioLAN Giant Anole • Schivartz
119
"albus" for "white" and "ocellus" for "eye,"
in allusion to the white spots that are typi-
cal of the holotype.
Anolis baleatus Cope
Eti])ristis baleatus Cope, 1864, Pvoc. Acad. Nat.
Sci. Philadelphia, p. 168.
Type locality. Santo Domingo; holo-
type, British Museum (Natural History)
1946.8.29.22.
Definition. A giant species of Hispanio-
lan Anolis characterized by the combina-
tion of large size (males to 1<S0 mm,
females to 148 mm snout-vent length),
snout scales at level of second canthal
scales 2 to 5 (modally 2 or 4, by popula-
tion) but usually 2 or 3 (75 percent), ver-
tical loreal rows 5 to 10 ( modes by popula-
tion 6, 7 or 8), scales between supraorbital
semicircles 1 to 4 (modally 3), interpari-
etal scales separated from supraorbital
semicircles modally by 4 or 5 scales, verti-
cal dorsal scales generally small ( 12 to 24
in standard-distance), ventral scales rela-
tively small ( 15 to 34 in standard-distance ) ,
nuchal crest scales in both sexes very high
to high, rarely moderate, never low, body
crest scales usually high to moderate, rarely
low, subocular scales usually not in contact
with supralabial scales; dorsal body colora-
tion and pattern usually some shade of
green, varying from dull greenish brown to
bright emerald green, either conspicuously
crossbanded with few (3 or 4) to very
many crossbands, in the latter condition the
lizards appearing tigroid, or, on the other
hand, without crossbanding but blotched,
never stiiped or with dark occipital, nu-
chal, or lateral dark markings, dewlap in
males from pale yellow to vivid orange, in
females from brownish or very pale yellow
to orange or gray, often suffused with gray-
ish or brownish, or nearly white, chin and
throat yellowish, green, or orange, often
with a dark dotted or mottled or reticulate
pattern, and pale subocular crescent absent
in adults.
Distribution. The eastern two-thirds of
the Repiiblica Dominicana, from Puerto
Plata, Santiago, and La Vega provinces
south to San Cristobal Province and the
Distrito Nacional, and east to La Altagracia
Province; also in and near the Sierra Mar-
tin Garcia and the southern slopes of the
Cordillera Central and the Sierra de Ocoa
in Azua and Peravia provinces; occurs on
Isla Saona but unrepresented by specimens
from that satellite island.
Anolis boleotus baleatus Cope
Type locality. "Santo Domingo"; here
restricted to the vicinity of Puerto Plata,
Puerto Plata Province, Repiiblica Domini-
cana (see rationale for this restriction be-
low ) .
Definition. A subspecies of A. baleatus
characterized by the combination of mod-
ally 4 snout scales between second canthal
scales, 7 vertical rows of loreal scales, 3
scales between the supraorbital semicircles,
moderate number of vertical dorsal scales
(14-21; mean 17.5), high number of ven-
tral scales (19-34; mean 23.8), nuchal crest
scales very high (usually) to high or mod-
erate (rarely), body crest scales high
(rarely) to moderate (usually), subocular
scales always in contact with supralabial
scales, males from pale green or rich bluish
green to brown dorsally, with three bright
yellow to darker green or greenish brown
irregular crossbands, lower sides usually
bright yellow, females apparently with the
same body patterns and hues as the males
(see below), throat in males bright yellow
to bright orange, rarely mottled with
brown, dewlap in males always vivid to
brilliant orange, and the upper surfaces of
hindlimbs bluish green, conspicuously
barred with bright yellow.
Distribution. Known from the Cordil-
lera Septentrional and the northern coastal
plain of the Repiiblica Dominicana, from
Puerto Plata, Espaillat, and Santiago prov-
inces, but probably occurring elsewhere in
this range and to the north of it; specimens
from Los Bracitos, Duarte Province, should
also be included ( on geographical grounds )
with A. b. baleatus, since Los Bracitos lies
120 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
in the eastern extremity of the Cordillera locality of the name to the vicinity of a ma-
Septentrional, but the specimens are old jor city that is presumed to lie within the
and greatly discolored and I have not con- area to which I ascribe this boldly cross-
sidered them as pertaining to the nominate banded subspecies,
subspecies. The series of 15 A. b. haleatus shows the
Discussion. Eupristis haleatus Cope was following variation. The largest males
named from a single specimen from "Santo (ASFS V33558, ASFS V18123) have snout-
Domingo." I have examined the holotype, vent lengths of 148, and the largest fe-
collected by A. Salle, in the British Museum male ( MCZ 128380) has the same dimen-
( Natural History ) . Considering its length sion. These three lizards are all from the
of time in preservative, it is in excellent Cordillera Septentrional north of Puesto
condition and shows a striking pattern of Grande. Snout scales at level of the second
three bold pale body crossbands on a canthal vary between 2 and 4; the mode
darker dorsal ground color, contrastingly is 4 (eight specimens). The vertical loreal
banded hindlimbs and tail, and immaculate rows vary between 5 and 9, with a mode
throat. The specimen is a female, and, un- of 7 (six specimens). There are 3 scales
fortunately, I have only two adult females between the supraorbital semicircles in all
from the range ascribed above to A. h. ha- specimens. There are modally 5 scales be-
leatiis: both are without color data in life, tween the interparietal and the semicircles;
At least one of them (MCZ 57717) resem- 5 scales are involved in 63 percent of the
bles the pattern of the haleatus holotype combination; actual counts are 4/4 (1), 4/5
to a striking degree. (4), 5/5 (7), 5/6 (1), 6/6 (1), and 5/7
Through the courtesy of Ernest E. Wil- (1). Vertical dorsals range between 14 and
liams, I have a copy of a map prepared by 21 ( mean 17.5 ) , horizontal dorsals between
William J. Clench which shows the locali- 16 and 26 (19.7), and ventrals between 19
ties where A. Salle is known to have col- and 34 (mean 23.8). Of nine adult males,
lected. Considering the era of his travels six have the nuchal crest scales very high,
(the mid-1800's), Salle traveled widely two have these scales high, and one has
throughout the Repiiblica Dominicana, them moderate. Of three females, the nu-
from (in the north) Puerto Plata, Ponton, chal crest scales are very high in two and
Santiago, Moca, La Vega and Cotui, east high in one. The body crest scales are high
to Higiiey, Cabo Engafio and San Rafael in one male and moderate in eight males;
del Yuma, in the eastern interior to Hato in three females, the body crest scales are
Mayor and El Seibo, along the southern high in one and moderate in two. All spec-
coast from Santo Domingo to San Cristo- imens have the subocular scales in contact
bal, Bani, Azua, Barreras, and Barahona, with the supralabial scales,
and into the Valle de San Juan to the city Males are usually conspicuously cross-
of San Juan. He also ascended the south- banded. Specimens have been recorded as
ern slopes of the Cordillera Central near pale green with three irregular darker
San Jose de Ocoa. Although much of Sal- green crossbands, brown with three faint
le's Dominican travels was in territory of green-brown crossbands, or rich bluish
A. haleatus, he was also in the ranges of green with three bright yellow crossbands.
A. ricorcli and A. harahonae. The holotype. The lower sides are bright yellow (which
as V^illiams (1962: 2, footnote 1) pointed grades into a grayish venter), and this color
out, has elongate nuchal crest scales, and also occurs on the throat, which varies
there is no doubt that the name haleatus from yellowish to bright yellow or orange,
is applicable to some population that pos- occasionally mottled with brown. The dew-
sesses this character. Since Salle traveled lap is brightly colored; it has been recorded
within the range of the northern population as "vivid orange," "bright vivid orange,"
of A. healeatus, I have restricted the type "brilliant yellow-orange," and "very bright
HisPANiOLAN Giant Angle • Schtvmiz 121
orange." The upper siirfaee of the head is
reddish brown and the hindhnibs are green
to bhiish green,' barred with bright yellow.
In general, male A. b. haleatus are vividly
patterned and eolored lizards. I have eol-
lected no females myself and thus have no
notes on this sex from life; however, one
recently (1971) collected female ( MCZ
128380) still is dark green with several thin
vertical pale crossband remnants on the
sides and back, and another female ( MCZ
57717) is contrastingly patterned in dark
and pale green, the latter occurring as ver-
tical crossbands.
The series includes three subadults, with
snout-vent lengths between 73 and 83.
One of these (ASFS V33559; snout-vent
length 80) was medium brown dorsally
with a black postocular streak and an or-
range dewlap that was streaked with black
basally. None of the subadults as pre-
served shows any crossbanding or other
pattern elements. It is interesting that the
only Hispaniolan giant anole taken at night
sleeping in the brown phase is the above
mentioned subadult.
Remarks. All ASFS specimens collected
by myself and parties were secured at
night while the lizards were sleeping.
Typical situations are in gallery forest and
cafetales along mountain streams in the
Cordillera Septentrional. Favored sleep-
ing sites for these lizards in the region are
pendant and semi-pendant woody vines;
Fowler reported that one adult male se-
cured by him at night was not asleep and
was slowly ascending a tree trunk as
Fowler approached. It is possible that this
lizard had been disturbed by the bright
light from Fowler's flashlight or by unfa-
miliar movements and noises, since I doubt
that any of the Hispaniolan giant anoles
are normally active at night. However, all
these lizards waken quickly when dis-
turbed and unless promptly secured, grad-
ually wander away into the greenery and
are lost to view. One of the juveniles was
secured only 6 feet (1.8 meters) above the
ground, whereas one of the adults was
shot from a tree limb 35 feet ( 10.7 meters )
above a mountain stream. The specimen
from near Sosi'ia was taken in dense hard-
woods on a limestone substrate.
The altitudinal distribution of A. 1). ha-
leatus is from 1400 to 2200 feet ( 427 to 671
meters), but the ta.xon occurs much lower
than this, since the specimen from near
Sosua was in limestone hills near sea level.
Specimens examined. REPUBLICA DO-
MINICANA: Espaillat Province, 2km N Pu-
esto Grande, 1400 to 2200 feet (427 to 671
meters) (ASFS V18048, ASFS V33557-
59); 5 km N Puesto Grande (MCZ
128380); 11 km N Puesto Grande, 2100
feet (641 meters) (ASFS V18123, ASFS
V18292): Puerto Plata Province, 11 km SE
Sosua (ASFS V1717); Santiago Province,
Pena (MCZ 57713, MCZ 57715-19); no lo-
cality other than Santo Domingo — British
Museum (Natural History) 1946.9.28.22—
holotype of Eupristis haleatus.
Anolis haleatus multistruppus
new subspecies
Holotype. USNM 193975, an adult
male, from Guaigiii, 3 mi. (4.8 km) S La
Vega, La Vega Province, 300 feet (92 me-
ters), Repiiblica Dominicana, one of a se-
ries taken by Danny C. Fowler, Albert
Schwartz, and Bruce R. Sheplan on 9 No-
vember 1971. Original number ASFS
V33680.
Paratopes. ASFS V33681-86, MCZ
125612-15, CM 54107-12, same data as ho-
lotype; ASFS V18547-50, same locality as
holotype, J. R. Dennis, J. A. Rodgers, Jr.,
and A. Schwartz, 27 July 1969.
Definition. A subspecies of A. haleatus
characterized by the combination of mod-
ally 2 snout scales between second canthal
scales, 7 vertical rows of loreal scales, 3
scales between the supraorbital semicircles,
4/4 scales between the interparietal and the
supraorbital semicircles, high number of
vertical dorsal scales (14-24; mean 18.6),
moderate number of ventral scales ( 18-29;
mean 22.3), nuchal crest scales very high
to high (usually) to moderate (rarely),
body crest scales high (rarely) to moder-
ate (usually), subocular scales almost al-
122 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
ways separated from supralabial scales by
one row of scales, both sexes as adults re-
taining the complex juvenile pattern of
many fine dark green, green, and yellow
vertical bars, occasionally (in females)
bright pea green with three pale green
crossbars more prominent than any other
dorsal pattern elements, throat green to
yellow green, dewlap in males very pale
yellow to very pale peach, suffused basally
with pale gray, in females very pale yellow
to pale yellow, strongly suffused with pale
gray to entirely pale gray.
Distribution. Known only from the type
locality but presumed to occur on the
northern and probably eastern lower faces
of the Cordillera Central in proper habi-
tats; possibly extending as far west on the
northern face of this range as the Rio Bao
near Los Montones (see discussion below).
Description of holotype. An adult male
with a snout-vent length of 146 and a
tail length (broken) of 97; snout scales at
level of second canthals 2, 9 vertical rows
of loreal scales, 3 scales between the su-
praorbital semicircles, 4/4 scales between
the interparietal and the supraorbital semi-
circles, vertical dorsals 21, horizontal dor-
sals 25, ventrals 29, one row of scales be-
tween the suboculars and supralabials,
fourth toe lamellae on phalanges II and III
30, nuchal crest scales high, body crest
scales moderate; in life, dorsal body pat-
tern of many fine green, dark green, and
yellow crossbands, upper surface of head
grayish tan in contrast to the brighter dor-
sal colors, chin and throat very pale yellow
or yellow-green with no clearly delineated
darker green markings, and dewlap very
pale yellow, much suffused basally with
gray.
Variation. The series of 21 A. h. multi-
struppus is composed of eight males and 13
females. The largest male has a snout-vent
length of 146 and is the holotype. The
largest female (ASFS V33684) has a
snout-vent length of 136 and is a topo-
type. Snout scales at the level of the sec-
ond canthals range between 2 and 5; the
mode is 2 (15 specimens). The vertical
loreal rows vary between 6 and 9, with a
mode of 7 (nine specimens). There are 2
or 3 scales between the supraorbital semi-
circles (mode 3). There are modally 4
scales between the interparietal and the
supraorbital semicircles; 4 scales are in-
volved in 58 percent of the combinations;
actual counts are 3/3 (1), 3/4 (2), 4/4
(8), 4/5 (5), 5/5 (3), and 5/6 (1). Ver-
tical dorsals range between 14 and 24
(mean 18.6), horizontal dorsals between
17 and 25 (20.6), and ventrals between 18
and 29 (22.3). All three adult males have
the nuchal crest scales high, and of ten
females, three have these scales very high,
five have them high, and two have them
moderate. All three males have the body
crest scales moderate, whereas two females
have the body crest scales high, eight have
them moderate, and one has them low. In
all but one specimen (5 percent), the sub-
oculars are separated from the supralabials
by 1 scale.
Adults of both sexes retain the juvenile
multibanded pattern of dark greens, me-
dium greens, and yellow. One adult fe-
male was recorded as bright pea-green
with three pale green crossbands, which
are remnants of the hollowed yellow cen-
ters of the five or six dark brown to dark
green crossbands. In general, the total as-
pect of adults and juveniles is of a con-
trastingly tigroid lizard, the stripes varying
shades of greens, yellows, and ( in the dark
phase) browns. The upper surface of the
head is grayish tan in males and tannish
green in females, and the throat is un-
marked green to yellow-green. One of the
most striking features of A. b. multistrup-
pus is the faded dewlap coloration. In
males, the colors vary between very pale
yellow and very pale peach, basally suf-
fused with pale gray. In females, the dew-
lap is even more drab, with pale yellow
the basic color, but the gray suffusion may
be so extensive as to limit the yellow pig-
ment to the dewlap edge or to cause the
dewlap to be pale gray.
The type series includes seven juveniles
and subadults, with snout-vent lengths be-
HisPANioLAN Giant Angle • Schwartz
123
twecn 47 and 99. These prcstMit a imi-
fonii aspect of multiple dorsal bands as
described above, and even tlie largest of
the subadnlts clearly shows this condition.
In life, a small juvenile (snout-vent length
53) was recorded as pale gray with a
yellow-green head and about four reversed
chevrons between the neck and the hind-
limbs, these chevrons being the pale hol-
lowed remnants of the darker crossbands,
wliich, in this individual, arc obscure. The
small lizard also had a black postocular
line and a charcoal postangular smudge.
The juvenile and subadult dewlaps are
pale flesh to very pale yellow, somewhat
suffused basally with light to very dark
gray.
Comparisom. Meristically, inulfistrup-
pus differs from nominate haleatus in hav-
ing 2 (rather than 4) snout scales at the
level of the second canthals, 4/4 (rather
than 5/5) scales between the interparietal
and the supraorbital semicircles, and in
having slightly less ventrals (means 22.3
and 23.8). There is also a strong tendency
for both sexes of haleatus to have very
high nuchal crest scales, whereas these
scales are more often only high in midti-
struppus. It is in color and pattern that
these two subspecies differ most strikingly.
In the introduction to the present paper I
commented on my having collected speci-
mens from the Cordillera Septentrional and
Guaigiii on two succeeding days, and on
the color and pattern differences being at
once very apparent. The bright orange
throat and dewlap of haleatus contrast
quite obviously with the pale yellow to
gray dewlaps in multistruppus. The body
patterns of the two subspecies likewise are
quite different, with the finely and multi-
banded multistruppus in contrast to the ir-
regularly banded dorsum with only three
bands in ])aleatus.
Discussion. A. h. multistruppus is
known with certainty from only a single
locality, which lies at the foot of the Cor-
dillera Central at an elevation of 300 feet
(92 meters). The locality is unique in that
it represents an extensive stand of original
lowland forest in this region, hardwood
forest which abuts upon the lower pine-
clad slopes of the mountains. This locality,
Guaigiii, is separated from the known
range of A. h. haleatus by the Vallc de
Cibao, which here is a moderately arid
and broad valley presently very much un-
der cultivation. I have seen no specimens
from this intervening valley but surely the
lizards occur there, despite the cultivation.
One other specimen requires mention.
This is a subadult male (ASFS V33856)
with a snout-vent length of 55, from 3.4
mi. (5.4 km) SE Los Montones, Rio Bao,
1600 feet (488 meters). This locality is on
the northern slopes of the Cordillera Cen-
tral, some 45 kilometers to the west of
Guaigiii, but separated from Guaigiii by
intervening, moderately high spurs of the
Cordillera Central. The specimen was se-
cured by a local boy in an area of high-
canopied forest along the Rio Bao. A visit
by ourselves to this area at night yielded
no A. haleatus, despite exceptionally fine
conditions. The lizard in life was all green
except for a white preaxillary bar, and the
dewlap was dull brownish. This specimen
in no way resembles comparably sized ju-
venile multistruppus, in either color or pat-
tern. Its status remains uncertain.
To the east, multistruppus must come in
close contact or intergrade with the sub-
species that occurs throughout the north-
eastern portion of the Republica Domini-
cana; details of this contact will be
discussed under the account of the latter
subspecies. Likewise, to the south, multi-
struppus may come into contact with the
subspecies in the high Cordillera Central;
details of this association will be discussed
under the description of the Central sub-
species.
Remarks. All specimens of A. h. multi-
struppus were collected on two occasions,
while the lizards slept at night. Young in-
dividuals were taken from generally low
situations on shrubs and the lower
branches of trees, whereas adults were ob-
served sleeping in the higher canopy; the
total range of heights was between 5 feet
124 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
and 25 feet (1.5 and 7.6 meters). The Rio scales always high, subocular scales al-
Camu flows through the Guaigiii woods, most always separated from supralabial
and many individuals were taken from scales by one row of scales, both sexes
tree limbs that overhang the river. either marbled or blotched with varying
The name multistruppus is from the shades of greens or browns, or dark brown
Latin "multus" for "many" and "struppus" banded with dull cream, never with many
for "thong, strap," in allusion to the many fine crossbars, venter in males pale green,
dorsal crossbands in this subspecies. flecked with darker green, and male dew-
laps pale yellow-orange to orange, gray
Anolis baleatus sublimis new subspecies basally and marbled green anteriorly, fe-
Holotijpe. CM 54104, an adult male, male dewlaps irregularly yellow-orange
from 0.3 mi. (0.5 km) E El Rio, 3800 feet with brown spotting.
(1159 meters). La Vega Province, Repub- Distribution. The uplands of the Do-
lica Dominicana, taken by Richard Thomas minican Cordillera Central at elevations
on 26 June 1963. Original number ASPS between 2000 and 4000 feet, in the area
X8114. between El Rio, La Palma, and Jarabacoa.
Paratypes. ASFS X8558, 4 km SW El Description of holotype. An adult male
Rio, 4000 feet ( 1220 meters ) , La Vega with a snout-vent length of 143 and a tail
Province, Republica Dominicana, R. F. length of 167 (regenerated); snout scales
Klinikowski, 2 July 1963; USNM 62104-05, at level of second canthals 3, 7 vertical
El Rio, La Vega Province, Republica Do- rows of loreal scales, 2 scales between the
minicana, W. L. Abbott, 19 May 1919; supraorbital semicircles, 4/4 scales be-
AMNH 41294, El Rio, La Vega Province, tween the interparietal and the supraor-
Repiiblica Dominicana, G. K. Noble, 31 bital semicircles, vertical dorsals 20, hori-
August 1922; ASFS V18594, La Palma, 14 zontal dorsals 21, ventrals 29, one row of
km E El Rio, 3500 feet (1068 meters). Re- scales between the suboculars and supra-
publica Dominicana, J. A. Rodgers, Jr., 30 labials, fourth toe lamellae on phalanges II
July 1969; MCZ 107019-21, La Palma, 14 and III 31, nuchal crest scales and dorsal
km E El Rio, 3500 feet ( 1068 meters ) , La body crest scales high; in life, dorsum
Vega Province, Republica Dominicana, na- dark brown banded with dull cream, this
tive collectors for E. E. Williams and A. S. pattern extending onto the tail, eyeskin
Rand, 25-31 July 1968; MCZ 128397, La grayish with a pale yellow eyering, venter
Palma, 14 km E El Rio, 3500 feet ( 1068 pale green, flecked with darker green, chin
meters). La Vega Province, Republica Do- and throat concolor with and patterned
minicana, T. P. Webster and R. B. Huey, like venter, dewlap pale yellow-orange,
6 July 1971; ASFS V18363-69, 8 km W grayish basally and marbled with green an-
Jarabacoa, 2000 feet (610 meters). La teriorly.
Vega Province, Republica Dominicana, J. Variation. The series of 18 sublimis is
A. Rodgers, Jr., 19 July 1969. composed of nine males and nine females.
Definition. A subspecies of A. baleatus The largest male (USNM 62104) has a
characterized by the combination of mod- snout-vent length of 150, the largest
ally 2 snout scales between second canthal female (MCZ 107021) 141; the male is
scales, 7 vertical rows of loreal scales, 3 from El Rio, the female from La Palma.
scales between the supraorbital semicir- Snout scales at the level of the second
cles, 4/4 scales between the interparietal canthals range between 2 and 5; the mode
and the supraorbital semicircles, high num- is 2 (10 specimens ) . The vertical loreal
ber of vertical dorsal scales (17-21; mean rows vary between 6 and 9, with a mode of
19.2 ) , high number of ventral scales ( 19- 7 ( eight specimens ) . There are 2 to 4
32; mean 25.1), nuchal crest scales very scales between the supraorbital semicir-
high (usually) to high (rarely), body crest cles (mode 3). There are modally 4 scales
HisPANioLAN Giant Angle • Schwartz 125
between the interparietal and the supraor-
bital semieircles; 4 scales are involved in
65 percent of the combination; the actual
counts are 3/3 ( 1), 3/4 ( 1), 4/4 ( 10), 4/5
(1), 5/5 (3), and 5/6 (1). Vertical dor-
sals range between 17 and 21 (mean 19.2),
horizontal dorsals between 17 and 24
(20.4), and ventrals between 19 and 32
(25.1). Of the six adult males, three have
the nuchal crest scales very high and three
ha\e them high, whereas all five adult fe-
males have these scales very high. All
adults of both sexes have the dorsal body
crest scales high. Three lizards ( 17 per-
cent) have the subocular scales in contact
with the supralabials.
In the green phase, adults of both sexes
are irregularly marbled or blotched with
varying shades of green or browns,
whereas in the brown phase, the body is
dark brown with three cream crossbands.
In males the venter and the chin and
throat are pale green, flecked or mottled
with darker green, the flecking or mottling
variably expressed in the series. The dew-
lap in males is pale yellow-orange to or-
ange, gray basally and often with marbled
green markings anteriorly, these markings
being a continuation of the dark green
throat markings. In females, the dewlap is
irregularly mottled with yellow-orange and
has some brown spotting. As preserved,
the series is remarkably uniform in show-
ing vague pale-and-dark marblings or mot-
tlings, and no adult shows any indication
of crossbands.
The series of paratypes includes six ju-
veniles and subadults, with snout-vent
lengths between 49 and 94. The three
smallest of these (49-70) are presently
patternless, as is also a specimen with a
snout-vent length of 73. Two other sub-
adults (snout-vent lengths 75 and 94)
show vague indications of mottled dorsum
with (in the larger) three slightly paler
dorsal crossbands. The larger of these two
specimens was recorded in life as dark
green dorsally with pale green crossbands,
and the interbars are mottled or marbled
with greens. The smallest juvenile noted
above was bright yellow-green in life and
had the venter slightly paler yellow-green;
the concealed surfaces of the thighs were
lead-gray, bordered above by bufl^y. The
absence of pale crossbars in very young
specimens of S'tihli)nis is noteworthy
Comparisons. There are no meristic
counts that separate sul)Iimis from adja-
cent imiltisfruppus; the means of ventral
scales in the two subspecies difl^er slightly,
however (22.3 in sul)li77iis, 25.1 in multi-
struppus). There is also a tendency for
su])limis to have more consistently very
high to high nuchal crest scales. The two
subspecies diff^er abundantly in body pat-
tern, however, with miilfistruppus having
many fine dorsal crossbands and sublimis
having basically a blotched dorsal pattern
with three bars present in some instances.
The juveniles of these two subspecies are
equally as distinct as the adults are in dor-
sal body pattern. The dewlaps are brighter
in male su])limis than in male muUistnip-
pus, the latter tending toward pale yellow
and yellow-grays, whereas in the former
the dewlaps are yellow-orange to orange,
although there is a gray basal suff^usion.
The ventral and throat flecking or mot-
tling in siihlimis differs from the unmarked
condition in mtiltistruppiis.
A. h. sublimis differs from A. h. hcileatiis
in having 2 (rather than 4) snout scales at
the level of the second canthals, 4/4
(rather than 5/5) scales between the in-
terparietal and supraorbital semicircles,
higher means of vertical dorsal scales ( 19.2
versus 17.5) and ventrals (25.1 versus
23.8). Both subspecies have very high to
high nuchal crest scales. In color, haleaius
is much the brighter, with an immaculate
bright yellow to orange throat and bright
yellow to orange dewlap in males, whereas
the dewlaps in sublimis are as bright as
those in haleatus but have a gray basal
wash. The patterned throat and venter in
sublimis differ from the immaculate throat
in buleatus. The dorsal patterns of these
two subspecies likewise are quite different,
that of baleatus regularly consisting of
126 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
three pale crossbands, whereas that of sub-
limis is mottled or blotched.
Discussion. A. h. suhlimis is closest
geographically to miiltistruppus; the two
subspecies are known from localities sep-
arated by only 20 kilometers airline
(Guaigi^ii and 8 km W Jarabacoa), but
minimally by a 1700-foot (519 meters) dif-
ference in elevation and by extensive
stands of pine forest, a habitat which no
Hispaniolan giant anole occupies. All spec-
imens of suhlimis were collected in mon-
tane gallery forest along streams, and the
subspecies appears to be restricted to this
sort of situation. Rand and Williams
(1969: 9) noted that they collected one
juvenile about 10 feet (3 meters) up on a
small branch of a forest tree at La Palma,
and that two adults were brought to them
by natives from a large tree in a nearby
agricultural area. A. h. suhlimis is thus
not known to come into contact with multi-
struppus on the northern slopes of the Cor-
dillera Central nor with the yet-to-be-de-
scribed subspecies to the east in the
Dominican lowlands. Likewise, it should
be recalled that the southwestern slopes of
this range are occupied by A. r. ricordi;
the nearest localities for ricordi and sul)-
lirnis (Juan de Herrera; south of El Rio)
are separated by about 70 kilometers air-
line, but this intei^vening area is composed
of the rugged and very high massif of the
Cordillera Central whose upper elevations
are covered by pine. It seems unlikely that
ricordi and suhlimis come into contact di-
rectly across the Cordillera.
The juvenile (ASFS 33856) from near
Los Montones upon which I commented
in the discussion of A. h. miiltistruppus
may be correctly assigned to suhlimis, since
the habitat and elevation for that specimen
is much more like that for suhlimis than
multistruppus. In color and lack of pattern
it agrees quite well with small suhlimis,
but until adults have been collected in the
Los Montones region (which lies some 30
kilometers to the northwest of Jarabacoa,
the closest suhlim.is locality) I am reluc-
tant to extend the known range of suhlim,is
into that area. It is this Los Montones A.
haleatus which is closest geographically
(50 kilometers) to an A. ricordi locality
( Los Quemados ) in the northwestern por-
tion of the Republica Dominicana.
Remarks. All ASFS A. h. suhlimis were
taken at night while asleep. All situations,
as noted above, were stream-associated
hardwood forest and cafetales, and the liz-
ards slept on vines and branches in their
customary fashion. The restriction of suh-
limis to riverine gallery forest is doubtless
artificial, since it is only along rivers and
streams in this area that any of the original
montane hardwood forests still remain. In
one case (west of Jarabacoa) the stream
was extremely steep, whereas in others the
streams were level. The altitudinal distri-
bution ( to which the name suhlimis refers )
is high. Only A. r. viculus reaches as high
an elevation in the Massif de la Hotte in
southwestern Haiti.
Anolis baleofus caeruleolatus
new subspecies
Holotype. USNM 193976, an adult
male, from 1.0 mi. (1.6 km) S Caiio Abajo,
>/Iaria Trinidad Sanchez Province, Repub-
lica Dominicana, one of a series collected
by native collectors on 28 November 1971.
Original number ASFS V34486.
Paratypes. CM 54119-26, MCZ 125628-
33, ASFS V34502-13, same data as holo-
type; AMNH 6017, Villa Riva, Duarte
Province, Republica Dominicana, C. R.
Halter, May-July 1915.
Associated specimens. REPCBLICA
DOMINICANA: Duarte Province, Los
Bracitos (AMNH 41465-66); ca. 4 km NE
Ponton (Rio Cuaba) (ASFS V2987); San-
chez Ramirez Province, 1 km SE La Mata
(ASFS V33650-51); La Vega Province,
12.8 km NW Bonao, 1200 feet (366 meters)
(ASFS V4317); 71 km NW Santo Domingo
(= near La Cumbre) (MCZ 128369); San
Cristohal Province, 5.0 mi. (8.0 km) NE
Gonzalo, 1000 feet (305 meters) (ASFS
V29420-21).
Definition. A subspecies of A. haleatus
characterized by the combination of mod-
HisPANioLAN Giant Anole • Schwartz
127
ally 4 scales between second canthal scales,
8 vertical rows of loreal scales, 3 scales be-
tween the supraorbital semicircles, 5/5
scales between the interparietal and the
supraorbital semicircles, moderate number
of vertical dorsal scales (14-22; mean
17.1), moderate number of ventral scales
(15-32; mean 22.4), nuchal crest scales
very high to high (usually) to moderate
or even low (rarely) in both sexes, body
crest scales extremely variable, modally
moderate in both sexes, but with some oc-
cinrences of high and many occurrences
of low body crest scales, subocular scales
almost always separated from supralabial
scales by one row of scales, both sexes
some shade of green (usually dark) with
foiu- pale green crossbars and with bright
sky-bhie blotches along the junction of the
green dorsal color and the paler venter
(less prominent in females than in males),
dewlap in males pale yellow to orange, in
females pale yellow to orange but with
much dark brown to grayish streaking or
smudging, throat in males deep yellow-or-
ange and immaculate or with very faint
greenish dots, in females yellow-green to
bright yellow, always with some darker
green dots, rarely marbled with dark green,
but never streaked with that color.
Distribution. Northeastern Republica
Dominicana, from Duarte, Sanchez Rami-
rez, La Vega, and northern and eastern
San Cristobal provinces, to the base of the
Peninsula de Samana (Caiio Abajo); in-
tergrades with the subspecies to the south
and east in the region of El Seibo Province.
Description of holotijpe. An adult male
with a snout-vent length of 137 and a tail
length of 250; snout scales between second
canthals 4, 7 vertical rows of loreal scales,
3 scales between the supraorbital semicir-
cles, 6/6 scales between the interparietal
and supraorbital semicircles, vertical dor-
sals 16, horizontal dorsals 23, ventrals 26,
one row of scales between the suboculars
and supralabials, fourth toe lamellae on
phalanges II and III 30, nuchal crest scales
very high, body crest scales moderate; in
life, dorsum dark green with four pale
green crossbars, the dark green color
blending fjuickly at the junction of the dor-
sal and ventral color into a series of diag-
onally directed sky-blue areas that give a
ragged appearance to the jmiction of the
dorsal and ventral colors; dorsal crossbands
continue onto the tail; cascjue gray-green,
eyeskin pale pea-green; dewlap pale yel-
low-orange, chin slightly deeper yellow-or-
ange, throat yellow-orange, immaculate
except for some vague pale greenish
smudges posterolaterally.
Variation. The series of A. b. caeruleo-
lattis consists of 20 males and 17 females.
The largest male (ASFS V34505) has a
snout-vent length of 148, the largest fe-
male (AMNh' 6017) 145. The male is
from the type locality, the female from
Villa Riva. Snout scales at the level of the
second canthals range between 2 and 5;
the mode is 4 ( 14 specimens ) . The verti-
cal loreal rows vary between 6 and 10; the
mode is 8 ( 15 specimens ) . There are 2 or
3 scales between the supraorbital semicir-
cles (mode 3). There are modally 5 scales
between the interparietal and the supraor-
bital semicircles; 5 scales are involved in
52 percent of the combinations; actual
counts are 4/4 (3), 4/5 (6), 5/5 (10), 5/6
(7), 6/6 (4), 6/7 (1), 4/6 (1), and 5/7
( 1 ) . Vertical dorsals range between 14
and 22 (mean 17.1), horizontal dorsals be-
tween 15 and 25 ( 19.9 ) , and ventrals be-
tween 15 and 32 ( 22.4 ) . Of 16 adult males,
four have the nuchal crest scales very high,
11 have them high, and one has them mod-
erate. Of 17 adult females, four have the
nuchal crest scales very high, ten have
them high, and three have them moderate.
In the adult males, the body crest scales
are high in six males, moderate in eight,
and low in two, whereas in the adult fe-
males, these scales are high in five, mod-
erate in six, and low in six. All but two
lizards (6 percent) have the suboculars
separated by one row of scales from the
supralabials.
In a series of 12 adult male topotypes,
the dorsal ground color was recorded as
some shade of green (usually dark green)
128 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
with four pale pea-green crossbands. The patterned hke adults except that the sky-
dorsal green color blends quickly ventro- blue lower edges to the dorsal color were
laterally into a series of irregular sky-blue absent and the dewlap was streaked brown
patches or blotches that mark the border and gray basally. The chin and throat
between the dorsal green and the pale yel- were immaculate pale green. There are no
low to cream venter. These sky-blue color data on the other juveniles, and none
patches are often prominently extended of them presently shows any pattern,
onto the lateral margins of the venter as a Comparisons. A. h. caeruleolatus dif-
series of diagonal, posteriorly directed fers from all previously described subspe-
areas, which, upon preservation, are still cies in».having the sky-blug patching along
prominent features of the lower sides. The the lower sides. In having four dorsal pale
upper surface of the head was gray-green green body bands, caeruleolatus differs
to brown, the eyeskin pale pea-green. The strikingly from multistruppus with its mul-
dorsal banded pattern of dark and light tiple banding; in addition, the dewlap of
green continues onto the tail. The dewlap multistruppus is pale and often grayish, in
is pale yellow-orange, yellow, or orange, contrast to the generally brighter dewlaps
and the chin is slightly deeper yellow-or- of caeruleolatus. From nominate baleatus,
ange, concolor with the throat, which is caeruleolatus differs in having the throat
either immaculate (usually) or with very yellow to yellow-green rather than bright
faint greenish dots or smudges. Eleven fe- yellow to orange, and female caeruleolatus
male topotypes were colored and patterned have the throat with dark green markings,
dorsally like the males, with the pattern From high upland sublitnis, caeruleolatus
extending onto the tail, but there is only a differs in having the sky-blue blotches ven-
vague indication of the ventrolateral sky- trolaterally and in lacking ventral mark-
blue pigmentation. The necks of females ings, and whereas caeruleolatus has com-
were often streaked with dark and pale parably pigmented dewlaps^ those in
greens. The chin and throat were yellow sublimis are generally paler and often suf-
to yellow-green, regularly with some fus^d at least basally with gray. The dor-
darker green dots, blotches, or occasionally sal patterns of both sublimis and caeruleo-
marbled with dark green. The female dew- latus are comparable, since both are
lap was yellow to pale orange, streaked crossbanded.
with dark brown or grayish. As far as meristic counts are concerned,
Two females from the haitises region caeruleolatus differs from the named sub-
near Gonzalo were deep to emerald green species in the following ways. Compared
in life with yellow dewlaps having varying with baleatus, caeruleolatus has modally 8
amounts of brown streaking or smudging; (rather than 7) vertical loreal rows, and a
the limbs were contrastingly banded dark lower mean number of ventral scales (22.4
and pale green. The throats were bright versus 23.8). There is also a strong ten-
yellow to bright green, with scattered dency for adult caeruleolatus to have mod-
deeper green spots in each case. In a pair erate to low body crest scales, whereas in
from La Mata, the dorsa were bright baleatus the tendency is toward high to
green, somewhat marbled with yellow and moderate body crest scales. Compared
yellow-green, the upper surfaces of the with multistruppus, caeruleolatus has mod-
heads were pale fawn, the eyeskin pale ally 4 (rather than 2) snout scales at the
grayish green, and the dewlaps orange in level of the second canthals, 8 rather than
both sexes. 7 vertical rows of loreals, 5/5 rather than
The series of A. b. caeruleolatus includes 4/4 scales between the interparietal and
four juveniles and subadults with snout- the supraorbital semicircles, and a lower
vent lengths from 60 to 91; the largest of mean of vertical dorsal scales (17.1 versus
these is a topotype that was colored and 18.6). With regard to body crest scales,
HisPANioLAN Giant Angle • Schwartz 129
these two subspecies show the same situa-
tion as caeruleolatus and baleatus. Com-
pared with .suJ)limis, caeruleolatus has 4
(rather than 2) snout scales at the level of
the second canthals, 8 (rather than 7) ver-
tical rows of loreals, 5/5 (rather than 4/4)
scales between the interparietal and the
supraorbital semicircles, and lower means
of both vertical dorsals (17.1 versus 19.2)
and ventrals (22.4 versus 25.1). A. h. suh-
limis has not been recorded as having the
dorsal body crest scales other than high,
in contrast to the strong tendency in cae-
ruleolatus of having these scales moderate
to low.
Discussion. A. h. caeruleolatus centers
in the extremely mesic eastern portion of
the Valle de Cibao in that area that has
the most rainfall in the Republica Domin-
icana. I have already commented on the
specimens from Los Bracitos, Duarte Prov-
ince; these specimens are old and pattern-
less and are from a locality in the Cordillera
Septentrional which is, farther west, occu-
pied by A. h. baleatus; I include them with
caeruleolatus provisionally. The specimen
from Ponton, Duarte Province, is a juve-
nile (ASFS V2987; snout-vent 60) and is
presently patternless; no color data are
available. It too I only provisionally re-
gard as caeruleolatus. The two specimens
from La Vega Province (ASFS V4317,
MCZ 128379) are also without color data
in life, and the former is a patternless ju-
venile (snout-vent 69). Specimens from
these last two localities also require verifi-
cation as to subspecfic status.
A. h. caeruleolatus presumably inter-
grades with four subspecies: baleatus, mul-
tistruppus, the subspecies on the Peninsula
de Samana, and subspecies to the south-
east. Only in the last case are specimens
that I interpret as intergradient known,
and they will be discussed under the de-
scription of the southeastern subspecies.
No intergrades are known between the
Samana subspecies, baleatus, or multistrup-
pus. Distance between caeruleolatus and
the nearest localities for these subspecies
are: Samana subspecies — 13 kilometers
(Caiio Abajo and 5 km NW Sanchez);
baleatus — 50 kilometers (Los Bracitos and
Pena); multistruppus — 12 kilometers (12.8
km NW Bonao and Guaigiii). Of these
presumed areas of contact, that between
caeruleolatus and the Samana subspecies is
not unexpected; the area between the two
known localities is very open and relatively
barren and devoid of trees and appears al-
ways to have been so. There are fine high
swamp-forests in the western part of this
intervening region, and it is possible that
intergrades between these two distinctive
subspecies will be encountered in these
forests. Most puzzling is the absence of
intergradation between caeruleolatus and
multistruppus. The specimen from north-
west of Bonao is a juvenile, but it does not
show the characteristic multiple crossbands
of both young and adult multistruppus. It
may be that multistruppus occupies only
the foothills of the Cordillera Central and
that the zone of intergradation between
multistruppus and caeruleolatus is very
abrupt.
Remarks. A. b. caeruleolatus is known
from sea level to an elevation of 1000 feet
(305 meters) in the haitises region near
Gonzalo and 1200 feet ( 366 meters ) north-
west of Bonao. Specimens were secured
primarily from native collectors; the long
series of topotypes is due to the industry
of the inhabitants of Cafio Abajo. The
Cafio Abajo area is one of cafetales and
cacaotales with high canopied shade-trees,
and the lizards apparently are extremely
abundant in this optimal habitat. The
pair of lizards from La Mata were secured
by me while they were copulating on the
side of a large shade-tree in a cafetal about
4 feet (1.2 meters) above the ground at
1225 hours. The two females from Gon-
zalo were taken during the day on large
trees adjacent to a small spring in the
haitises; the surrounding area was under
heavy cultivation, but the doline slopes
were covered locally with undisturbed for-
est.
The name caeruleolatus is from the
Latin "caeruleus" for "blue" and "latus"
130 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
for "side," in allusion to the sky-blue lower
sides of this subspecies.
Anolis baleatus samanae
new subspecies
Holotype. CM 54105, an adult male,
from 7.6 mi. (12.2 km) NE Sanchez, 1000
feet (305 meters), Samana Province, Re-
publica Dominicana, one of a series col-
lected by native collectors on 28 November
1971. Original number ASFS V34474 .
Paratypes. ASFS V34475-79, same data
as holotype; USNM 193990-92, same local-
ity as holotype, native collectors, 27 No-
vember 1971; MCZ 125634, 5.0 mi. (8.0
km) NW Sanchez Province, Republica Do-
minicana, J. Aria, 27 November 1971; ASFS
V34495-96, 5.0 mi. (8.0 km) NW Sanchez,
Samana Province, Republica Dominicana,
J. Aria, 28 November 1971; CM 54127-30,
5.0 mi. (8.0 km) NW Sanchez, Samana
Province, Republica Dominicana, J. Aria,
30 November 1971; MCZ 12563.5-39,
USNM 193993-4001, 5.0 mi. (8.0 km) NW
Sanchez, Samana Province, Republica Do-
minicana, J. Aria, 1 December 1971; ASFS
V34514, ASFS V34836-38, Las Terrenas,
Samana Province, Republica Dominicana,
native collector, 28 November 1971; ASFS
V1904, 6 km E Sanchez, Samana Province,
Republica Dominicana, R. Thomas, 30 Oc-
tober 1963; AMNH 28651, Samana, Sa-
mana Province, Republica Dominicana, J.
King, August 1924; AMNH 39817-23,
AMNH 42285, Laguna, Samana Province,
Republica Dominicana, W. G. Hassler, Oc-
tober-December 1929; USNM 61928, Cayo
Hondo, Samana Province, Republica Do-
minicana, W. L. Abbott, February 1919.
Definition. A subspecies of A. baleatus
characterized by the combination of mod-
ally 2 snout scales at level of second can-
thai scales, 7 vertical rows of loreal scales,
3 scales between the interorbital semicir-
cles, 4/4 scales between the inteiparietal
and the supraorbital semicircles, moderate
number of vertical dorsal scales ( 13-20;
mean 16.6), moderate number of ventral
scales (16-29; mean 22.1), nuchal crest
scales very high to high (usually) to mod-
erate or low (rarely) in both sexes, body
crest scales high to moderate but often low
in both sexes, subocular scales almost al-
ways separated from supralabial scales by
one (rarely 2) row of scales; dorsum in
both sexes in life blotched dark green,
greenish, dull gray-green, brown, or black-
ish, dewlaps in males dull yellow to pale
yellowish orange, in females very pale yel-
low to pale yellowish orange, streaked with
blackish or brown basally, and chin and
throat in males cream to yellowish or yel-
low-orange, mottled with black or gray, in
females pale green to greenish yellow with
dark green to brown streaking or even re-
ticulate.
Distribution. The Peninsula de Samana
in the northeastern Republica Dominicana,
and apparently islets in the Bahia de Sa-
mana.
Description of holotype. An adult male
with a snout-vent length of 145 and a tail
length of 222 (regenerated); snout scales
between second canthals 3, 6 vertical rows
of loreal scales, 3 scales between the supra-
orbital semicircles, 4/4 scales between the
interparietal and the supraorbital semicir-
cles, vertical dorsals 18, horizontal dorsals
19, ventrals 21, one row of scales between
the suboculars and supralabials, fourth toe
lamellae on phalanges II and III 30, nuchal
crest scales very high, body crest scales
high; in life, dorsum mottled dull greens
and gray-brown with whitish (almost
cream but suffused with pale gray); upper
surface of head mixed dark brown and
gray, venter dull greenish, dewlap orange,
chin and throat creamy to yellowish, not
marked with green.
Variation. The series of 54 A. b. sama-
nae consists of 32 males and 22 females.
The largest male (AMNH 39807) has a
snout-vent length of 157; the largest fe-
males (CM 54130, USNM 193994) have
snout-vent lengths of 145. The male is
from Laguna, the females from 5.0 mi. NW
Sanchez. Snout scales at the level of the
second canthals range between 2 and 5;
the mode is 2 (24 specimens). The verti-
cal loreal rows vary between 5 and 9; the
HisPANioLAN Giant Angle • Scliwartz 131
mode is 7 (25 specimens). There are 2 or
3 scales between the supraorbital semicir-
cles (mode 3). There are modally 4 scales
between the interparietal and the supraor-
bital semicircles; 4 scales are involved in
43 percent of the combinations: actual
counts are 3/3 (2), 3/4 (3), 4/4 ( 17), 4/5
(8), 5/5 (13), 5/6 (5), 6/6 (1), 6/7 (2),
4/6 (1), and 3/5 (1). Vertical dorsals
range between 13 and 20 (mean 16.6),
horizontal dorsals between 13 and 27
(19.3), and ventrals 16-29 (22.1). Of 30
adult males, 14 have the nuchal crest scales
very high, 15 have them high, and one has
them moderate; in 20 adult females, nine
have the nuchal crest scales very high,
nine have them high, one has them mod-
erate, and one has them low. Body crest
scales in males are high in three lizards,
moderate in 16, and low in ten; in females,
11 have these scales moderate and ten have
them low. The suboculars are separated
from the supralabials by one row of scales
in all but four specimens (7 percent),
which have them in contact, and one spec-
imen (2 percent), which has 2 rows of
scales in this position.
A. ]). samanae is basically a blotched liz-
ard, and no adults show any indication of
crossbanding. The body is irregularly
blotched with blackish, dark green, dull
green, gray-brown, and occasionally there
are sky-blue areas along the ventrolateral
margin of the dorsal coloration in males,
but these areas are not so prominent as in
caeruleolatus. Regardless of the dorsal
shades, the upper surface of the head is
mixed dark brown and shades of gray in
both sexes. The hindlimbs are finely
barred with pale and dark green. The
venter is dull greenish in both sexes. The
dewlap in males varies from dull yellow or
pale yellowish orange to orange, and the
chin and throat are yellowish, cream, or
yellow-orange, mottled with black or gray.
In females, the dewlaps are very pale yel-
low, pale yellow-orange, or grayish orange,
at times streaked with blackish or brown
basally, and the chin and throat ground
color is pale green, marbled, streaked, or
even reticulate with dark green to (rarely)
brown.
There arc one juvenile (AMNH 28651;
snout-vent length 40) and two subadult
(snout-vent lengths 92 and 97) A. 1). sa-
manae. The subadults are old and discolored
but their patterns seem not to differ from
those of full adults. The juvenile on the
other hand, has fom- bold pale crossbars on
the dorsum, the pattern continuing onto
the tail. This young individual has the
umbilicus still present and is presumably
near hatchling size.
Comparisons. Since samanae and cae-
ruleolatus are adjacent geographically, the
most pertinent comparisons are between
them. Examples of these two populations,
as noted in the introduction to the present
paper, were available to me simultaneously
and I was struck with their differences in
life. A. 1). samanae is a blotched lizard
whereas caeruleolatus is a crossbanded
one; the latter subspecies also typically has
sky-blue ventrolateral blotches, a feature
absent (or occasionally poorly expressed)
in male samanae. Male dewlap colors are
similar in both subspecies, although fe-
male dewlap colors in samanae seem some-
what paler than those of caeruleolatus.
The chin and throat markings of the two
subspecies are quite distinct; in male cae-
ruleolatus, the throat is deep yellow to
yellow-orange, at best with very faint gray-
ish dots or smudges, whereas in male
samanae the throat is yellowish or cream
to yellow-orange, mottled with black or
gray. In female caeruleolatus, the throat
is yellow to yellow-green, always with
some dark green dots, blotching, or mar-
bling, whereas in samanae females, the
throat is pale green, greenish yellow, or
yellow-green, with dark green to brown
streaking or reticulum.
The only subspecies thus far described
which is blotched like samanae is the Cor-
dillera Central suhlitnis, although caerul-
eolatus may show a marbled dorsum in
some areas. No pigmental or pattern dif-
ferences separate samanae and sublimis,
since in both dorsal coloration and color of
132 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
the dewlap the major color involved is
green. However, the throat in male sub-
limis is pale green, whereas in samanae it
is cream to yellow-orange. Certainly rnulti-
struppus and samanae are easily distin-
guished in the field by their very different
dorsal patterns, for example, and haleatus,
with its very, very bright chin and thioat,
both of which are immaculate, is quite dis-
tinctive from samanae.
In meristic data, samanae differs from
caeruleolatus in having 2 (rather than 4)
snout scales, 7 (rather than 8) vertical
rows of loreals, and 4/4 (rather than 5/5)
scales between the inteiparietal and the
supraorbital semicircles. From multistrup-
pus, samanae differs in having a lower
mean of vertical dorsal scales ( 16.6 versus
1S.6), and the same difference occurs be-
tween samanae and su])Umis ( 16.6 versus
19.2) and in ventrals (22.1 versus 25.1).
From haleatus, samaiuie differs in having
2 (rather than 4) snout scales, 4/4 (rather
than 5/5) scales between the inteiparietal
and the supraorbital semicircles, and lower
means in both vertical dorsals ( 16.6 versus
17.5) and ventrals (22.1 versus 23.8). The
nuchal crest scales in samanae are more
consistently very high to high than they
are in any of the other subspecies of A.
haleatus.
Discussion. As pointed out in the dis-
cussion of A. h. caeruleolatus, there are no
intergrades known between that subspe-
cies and samanae. The isthmus of the Pe-
ninsula de Samana is much cleared and
locally even barren, but there are large
western swampy areas that support mag-
nificent hardwood forests toward the land-
ward side. These forests may well support
intermediates between samanae and cae-
ruleolatus, or, because of their proximity
to the mainland, they may be inhabited by
caeruleolatus. Specimens from 5.0 mi. NW
Sanchez, that locality for samanae which
is closest to a known locality for caeruleo-
latus ( 18 kilometers ) , show no tendencies
toward the crossbanded condition of cae-
ruleolatus.
A. h. samanae is the only Hispaniolan
giant anole known by specimens from any
off-shore island or islet. The specimen
from Cayo Hondo, taken by W. L. Abbott,
constitutes this record, although I am un-
able to locate this islet. I assume it is one
of the archipelago within the Bahia de Sa-
mana.
Remarks. All but one A. h. samanae se-
cured by myself and parties were native-
collected. The exception is a lizard taken
by Richard Thomas, one of two seen on a
small tree and in a vine tangle in a steep
limestone ravine east of Sanchez. The area
of the type locality is in the uplands of the
Sierra de Samana on the road between
Sanchez and Las Terrenas. Thus newly
constructed road passes through superb
mesic high-canopied forest, and much of
the area is not yet seriously disturbed. Ob-
viously from the number of lizards secured
by natives in this region, A. /;. samanae is
common. The range is not high, with a
maximum elevation of 1673 feet (510 me-
ters) in Monte Las Caiiitas; this mountain
lies between Sanchez and Las Terrenas.
Specimens from Las Terrenas itself were
secured by natives from near-coastal mesic
cafetales and cacaotales, and lizards from
northwest of Sanchez were in similar situ-
ations.
Only three other reptiles (Diplo^lossus
sternurus alloeides Schwartz, Leiocephalus
personatus pyrrholaemus Schwartz, and
Dromicus parvifrons niger Dunn) are known
to have differentiated at the subspecific
level on the Peninsula de Samana. Sphaero-
clactylus clenchi Shreve and Sphaeroclac-
tijlus samanensis Cochran both occur there
and have as yet unnamed populations, one
of which in each case is limited to the pen-
insula. It is also of interest to note that in
Anolis clisticJius Cope, the Samana popula-
tion is identical to the population on the
soutliern shores of the Bahia de Samana
(ignigularis Mertens), but that the range of
this subspecies is interrupted at the head
of the Bahia de Samana by A. cl. domini-
censis Reinhardt and Liitken (see Schwartz,
1968: 280-81, for details).
HisPANioLAN Giant Angle • Sclucmiz 133
Anolis baleatus litorisilva new subspecies
Holotype. USNM 193977, an adult
male, from 1.2 km SSW Piinta Cana, La
Altagracia Province, Rcpviblica Domini-
cana, one of a series collected by Danny
C. Fowler and Bruce R. Sheplan, on 24
November 1971. Original number ASFS
V35095.
Paratypes. ASFS \'35096-100, same
data as holotvpe; CM 54113-14, MCZ
125616-17, 5.5 km SSW Punta Cana, La
Altagracia Province, Republica Domini-
cana, D. C. Fowler, 27 November 1971;
ASFS V29090, Juanillo, La Altagracia
Province, Republica Dominicana, native
collector, 24 July 1971; ASFS V961-62, 0.5
mi. NW Boca de Yuma, La Altagracia
Province, Republica Dominicana, R. F.
Klinikowski, R. Thomas, 2 September 1963;
ASFS VI 136, 2.5 km NW Boca de Yuma,
La Altagracia Province, Republica Domin-
icana, native collector, 4 September 1963;
ASFS V17573, 4 km NW Boca de Yuma,
La Altagracia Province, Republica Domi-
nicana, A. Schwartz, 13 June 1969; ASFS
V17616, 2 km NW Boca de Yuma, La Al-
tagracia Province, Republica Dominicana,
J. B. Strong, 15 June 1969.
Definition. A subspecies of A. I)(ileatus
characterized by the combination of 2 or 4
scales at level of the second canthal scales,
7 vertical rows of loreal scales, 3 scales be-
tween the interorbital semicircles, 4/5
scales between the interparietal and the
supraorbital semicircles, low number of
vertical dorsals (13-19; mean 15.9), low
number of ventral scales ( 18-26; mean
21.3), nuchal crest scales always very high
to high in both sexes, body crest scales
high (rarely) to moderate or low, suboc-
ular scales usually separated from supra-
labial scales by one row of scales; dorsum
in life varying from light blue-brown to
light greenish brown in males, dull brown
to olive-brown in females, blotched with
creamy to gray, dewlap in males bright
orange, brownish in females, and chin and
throat (including lips) bright orange in
males, pale yellow-green in females.
Distri])ution. Extreme eastern Repub-
lica Dominicana in La Altagracia Province,
from Punta Cana to the \'icinity of Boca de
Yinna.
Description of holotype. An adult male
with a snout-vent length of 136 and a tail
length of 183 (regenerated); snout scales
between sc^cond canthals 2; 6 vertical rows
of loreal scales, 3 scales between the supra-
orbital semicircles, 5/5 scales between the
interparietal and the supraorbital semicir-
cles, vertical dorsals 15, horizontal dorsals
16, ventrals 25, subocular scales in contact
witli the supralabial scales, fourth toe la-
mellae on phalanges II and III 31, nuchal
crest scales very high, dorsal body crest
scales high; in life, dorsum blotched light
blue-brown and light green-brown; venter
pale gray-green; chin, lips, and dewlap
bright orange.
Variation. The series of 16 A. h. litori-
silva is composed of six males and ten fe-
males. The largest male (MCZ 125616)
has a snout-vent length of 158, the largest
female (ASFS V961) 131. The male is
from 5.5 km SSW Punta Cana, the female
from 0.5 mi. NW Boca de Yuma. Snout
scales at the level of the second canthals
range between 2 and 5; there are t\vo
modes, 2 and 4, each with five individuals.
The vertical loreal rows vary betwe(Mi 6
and 9; the mode is 7 (nine specimens).
There are 2 to 4 scales between the supra-
orbital semicircles (mode 3). There are
modally 4/5 scales between the interpari-
etal and the supraorbital semicircles; 5
scales are involved with 59 percent of the
combinations; actual counts are 4/4 (4),
4/5 (6), 5/5 (5), and 5/6 (1). Vertical
dorsals range between 13 and 19 (mean
15.9), horizcmtal dorsals between 14 and
22 (18.5), and ventrals between 18 and 26
(21.3). Of four adult males, three have
the nuchal crest scales very high and one
has them high; of five adult females, two
have these scales very high and three have
them high. In the males, the body crest
scales are high in one and moderate in
three, and in the females, these scales are
134 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
moderate in two and low in three. The
siiboculars are separated from the siipra-
labials by one row of scales in all but one
specimen (6 percent).
A. h. litorisilva is essentially a blotched
lizard whose colors do not include bright
or even medium greens. The color notes
on the holotype apply equally well to the
other adult males — the dorsum is blotched
with bluish browns and light greenish
browns, without any clear greens, and
the blotching is often more pronounced on
the head than on the body. In females, the
dorsum is dull brown to olive-brown with
only occasional slight remnants of a lighter
green pattern on the head; the blotching
in the female involves creamy to gray pig-
mentation. The venter is pale gray-green
or whitish green in males, pale greenish
gray in females. The dewlap in all adult
males was recorded as bright orange, and
brownish in females. In males, the chin
(including the lips) is bright orange, and
pale yellow-green in females. The upper
surface of the head in males is blotched
like the body and is dark chocolate in fe-
males. In females, the upper surfaces of
the hindlimbs were recorded as olive-
brown, blotched with cream to gray like
the dorsum.
The series of A. h. litorisilva contains
seven juveniles and subadults (snout-vent
lengths 45 to 88). The smallest juvenile
(ASFS V17573, female) was bright green
in life with four pale buffy crossbands and
dark green shadow-bars between the cross-
bands; the tail was ringed cream and dark
gray, and the venter was pale green. The
dewlap was yellow-green and gray. A
slightly larger female (ASFS V17616) with
a snout-vent length of 57 was yellow-green
dorsally and without bands, the head was
brown; the eyeskin was green, and the
venter yellow-green. The tail was banded
black and yellow-green, and the dewlap
was mainly brown with the scale rows yel-
low-green. A still larger female (ASFS
V1136) with a snout-vent length of 67 was
green, faintly crossbarred with grayish
green, and there were charcoal smudges on
the neck. Two male subadults with snout-
vent lengths of 71 and 83 (ASFS V35099-
100) from the type locality were recorded
by Fowler as follows: "One with a strong
vertical banding pattern alternating brown-
green and white-gray, which extends from
tip of tail to the head where it becomes
slightly more diffuse; on the other, the dor-
sal groimd color is dull brown with rem-
nants of banding pattern only around head;
the ventral ground color of the first is gray-
green with brown mottling, the second is
dull gray-brown; in both juveniles, the
dewlap is orange-green and the chin and
lips are green." The largest subadult
(ASFS V29090) was patternless green
above, and the dewlap was orange with
charcoal stripes; the specimen is a female.
Comparisons. Because of its blotched
(rather than crossbanded) pattern, litori-
silva requires comparison with samanae
and suhlimis. The general effect of the
dorsa of all three subspecies is quite sim-
ilar, but samanae and suhlimis are much
the brighter lizards, with greens predomi-
nant in the dorsal pigmentation. On the
other hand, litorisilva is a much more drab
lizard, without clear greens in the adults,
the tendency being toward more sombre
hues, primarily shades of browns. From
all other described subspecies, litorisilva
differs in being blotched rather than cross-
banded and also in having much less
gaudy dorsal colors. In meristic counts,
litorisilva differs from the remaining sub-
species in the following ways. From cae-
ruleolattis, litorisilva differs in having 7
(rather than 8) vertical loreal rows, and
lower means of vertical dorsals (15.9 ver-
sus 17.1) and ventrals (21.3 versus 22.4).
From rnultistruppus, litorisilva differs in
lower means of vertical dorsals ( 15.9 ver-
sus 18.6) and ventrals (21.3 and 22.3).
From stihlitiiis, litorisilva differs in having
lower means of vertical dorsals ( 15.9 ver-
sus 19.2) and ventrals (21.3 versus 25.1).
From haleatus, litorisilva differs in having
lower means of vertical dorsals ( 15.9 ver-
sus 17.5) and ventrals (21.3 versus 23.8).
Meaningful comparisons of litorisilva with
HisPANiOLAN Giant Anole • ScJiwartz 135
other subspecies in counts of snout scales,
and scales between the interparietal and
the supraorbital semicircles, are impossible
since litorisilva has a bimodal condition in
the former (and the bimodes are 2 and 4,
those counts which occur singly as the
mode in the other subspecies) and has a
mode of 4/5 in the latter (whereas all
other species have either 4/4 or 5/5).
Considering the fairly large series of litori-
silva (16 specimens), these two "abnormal"
conditions are puzzling. At least in the
case of 4/5 counts, the absence of 3/3 or
3/4 counts in litorisilva suggests that this
subspecies tends toward a 5/5 count.
Discussion. A. /;. litorisilva appears to
be the extreme eastern isolate of the more
widespread A. haleatus stock. It occupies
semi-arid forests on and near the coast (as
at Juanillo and Punta Cana) and on the
limestone ridge behind Boca de Yuma.
Both situations are far more xeric than is
customary for A. haleatus, and the faded
nongreen coloration of the adults is doubt-
less a response to the dry and open to
dense forest conditions of this region.
Nevertheless, individuals are quite con-
spicuous at night as they sleep exposed. A.
b. litorisilva presumably comes into con-
tact with the subspecies to the north and
west (named below) but intergrades are
presently unknown; in the vicinity of Hig-
iiey (the closest locality for the adjacent
subspecies) the lizards are more brightly
colored and crossbanded and quite unlike
litorisilva.
Remarks. All but one specimen of li-
torisilva were collected by myself and par-
ties. Individuals were found sleeping in
primarily coastal forest (to which the
name, from "litus" for "shore" and "silva"
for "forest," refers in Latin) at elevations
from 4 to 15 feet (1.2 to 4.6 meters) above
the ground. Generally, juveniles sleep
closer to the groimd and in more dense
situations than adults. One juvenile was
taken from a roadside Acacia, a most un-
usual situation (since Acacia is a distinct
xerophyte) for any giant anole. Several
adults were taken in dense viny tangles,
sleeping on the woody vines; the advan-
tage of this situation was made (piite ob-
vious wIkmi I attempted to catch a large
adult at night by hand. The light from my
flashlight wakened the lizard almost im-
mediately, and although 1 was extremely
careful not to jar any of the vines, this was
a vain endeavor. At the first jostling, the
lizard jumped to the ground and escaped
in the dry leaf litter and understory.
Anol'is haleatus scelestus new subspecies
Holotype. CM 54106, an adult male,
from 5.1 mi. (8.2 km) E Santo Domingo
(from Rio Ozama), Distrito Nacional, Re-
publica Dominicana, one of three collected
by David C. Leber and Richard Thomas
on 18 June 1964. Original number ASFS
V2460.
Paratopes. ASFS V2461-62, same data as
holotype; MCZ 125618-27, 8.4 mi. (13.4
km) NE La Romana, 100 feet (31 meters).
La Romana Province, Republica Domini-
cana, B. R. Sheplan, 22 November 1971;
CM 54115-18, USNM 193981-89, 8.4 mi.
(13.4 km) NE La Romana, 100 feet (31
meters). La Romana Province, Republica
Dominicana, D. C. Fowler, A. Schwartz,
17 July 1971; MCZ 16321, La Romana, La
Romana Province, Republica Dominicana,
E. Leider, 1922; ASFS V29284-300, 0.2 mi.
(0.3 km) N Otra Banda, 350 feet (107 me-
ters). La Altagracia Province, Republica
Dominicana, D. C. Fowler, A. Schwartz,
26 July 1971; ASFS V21699-700, 1 km NE
Higiiey, La Altagracia Province, Republica
Dominicana, J. R. Dennis, R. Thomas, 16
August 1969; USNM 193979-80, 0.7 mi.
(1.1 km) W Higiiey, La Altagracia Prov-
ince, Republica Dominicana, R. Thomas,
29 August 1963; ASFS V1038, 1 mi. (1.6
km) W Higiiey, La Altagracia Province,
Republica Dominicana, R. Thomas, 3 Sep-
tember 1963; ASFS V28757, 15.5 mi. (24.8
km) E San Pedro de Macoris, Rio Cumay-
asa. La Romana Province, D. C. Fowler,
12 July 1971; ASFS V28910-16, 15.5 mi.
(24.8 km) E San Pedro de Macoris, Rio
Cumayasa, San Pedro de Macoris Province,
Republica Dominicana, D. C. Fowler, A.
136 Bulletin Musewyi of Comparative Zoology, Vol. 146, No. 2
Schwartz, 16 July 1971; ASFS V28847, 15.5
mi. (24.8 km) E San Pedro de Macoris, La
Romana Province, Repiiblica Dominicana,
A. Schwartz, 15 July 1971.
Associated specimens. REPUBLICA
DOMINICANA: La Altagracia Province, 1
km SE Las Lisas (ASFS V17434-35); San
Cristobal Province, 8 km N Yamasa, 200
feet (61 meters) (ASFS V28656).
Definition. A subspecies of A. haleatus
characterized by the combination of mod-
ally 2 scales at level of the second canthal
scales, 7 vertical rows of loreal scales, 3
scales between the supraorbital semicircles,
5/5 scales between the interparietal and
the supraorbital semicircles, low number of
vertical dorsals ( 12-20; mean 15.4 ) , low
number of ventral scales ( 17-28; mean
21.1), nuchal and body crest scales always
very high to high in both sexes, subocular
scales usually separated from supralabial
scales by one (occasionally two) row of
scales; dorsum in both sexes either green
with three pastel green crossbands or dark
green flecked with light green, cream with
some greenish to brownish green smudges,
dewlap in males deep yellow to deep or-
ange, streaked or smudged with dark
brown to charcoal, and throat in females
dark green marbled with yellow and pale
green (males unrecorded).
Distribution. Southeastern Republica
Dominicana, from the Sierra de Yamasa
and the vicinity of Santo Domingo in the
west, east to the region about Higiiey and
Las Lisas in La Altagracia Province.
Description of Jiolotype. An adult male
with a snout-vent length of 152 and a tail
length of 267; snout scales between second
canthals 4; 8 vertical rows of loreal scales,
2 scales between the supraorbital semicir-
cles, 4/5 scales between inteiparietal and
supraorbital semicircles, vertical dorsals
16, horizontal dorsals 16, ventrals 22, sub-
ocular scales separated from supralabial
scales by one row of scales, fourth toe la-
mellae on phalanges II and III 34, nuchal
crest scales high, body crest scales moder-
ate; in life, dorsum olive-green with six
pastel green crossbands, tail and venter
light green; dewlap dark yellow.
Variation. The series of 61 A. b. sceles-
ttis consists of 27 males and 34 females; a
large number of the specimens are juve-
niles and subadults. The largest male
(ASFS V29284) has a snout-vent length
of 180, the largest female (ASFS V29286)
147; both are from near Otra Banda. Snout
scales at the level of the second canthals
range between 2 and 4; the mode is 2 (32
specimens). The vertical loreal rows vary
between 5 and 8, with a mode of 7 (25
specimens). There are 1 to 4 scales be-
tween the supraorbital semicircles (mode
3). There are modally 5/5 scales between
the interparietal and the supraorbital semi-
circles; 5 scales are involved in 49 percent
of the combinations; actual counts are 3/4
(2), 4/4 (14), 4/5 (14), 5/5 (17), 5/6
(11), 6/6 (1) and 4/6 (1). Vertical dor-
sals range between 12 and 20 (mean 15.4),
horizontal dorsals between 15 and 25
(18.8), and ventrals between 17 and 28
(21.1). Of 11 adult males, nine have the
nuchal crest scales very high and two have
them high. Of 16 adult females, nine have
these scales very high and seven have them
high. Body crest scales in males are high
in two lizards, moderate in eight, and low
in three; in females, the body crest scales
are high in two, moderate in eight, and low
in six. Fifty-three specimens have the sub-
oculars separated from the supralabials by
one row of scales, whereas in four lizards
( 7 percent ) these scales are in contact, and
in two lizards (3 percent) they are sepa-
rated by tv/o rows of scales.
In general, both sexes of A. b. scelestiis
show a pattern of about six or seven fine
crossbands that are often obscured by dor-
sal blotching. Colors are shades of greens,
with brighter green the base color and the
blotching tending toward darker shades.
The crossbands are lighter pastel shades of
green, and in some lizards the dorsal
ground color is olivaceous. Another vari-
ant, which is somewhat more prevalent in
females, is an olive green to dark green
HisPANioLAN Giant Angle • Schwaiiz
137
dorsum, flocked with pale green. Two fe-
males from near Higiiey showed still an-
other style of body pattern and color, with
the dorsal ground color cream with some
dark green to brownish green snuidges,
and the neck with alternating pale blue
and charcoal markings, the pale blue mark-
ings persisting onto the cheeks. In males
the upper surface of the head is brown,
and in females it is mixed brown and
green, with the snout and supraocular
scales deep green in some lizards. In fe-
males, the chin and throat are dark green,
marbled with yellow and pale green. The
dewlap is rather \'ariable; in males it has
been recorded as dark yellow or deep yel-
low to orange or dark orange, whereas in
females the dewlap varies from yellow to
dark orange with dark brown, olivaceous,
or charcoal streaking, marbling, or smudg-
ing. Although there are no color notes in
life, in the preserved lizards the eyeskin is
regularly pale gray, and I presume that in
life the eyeskin is set off from the rest of
the head color in some pigmental fashion.
Many specimens of both sexes have the
lower sides tigroid with "stripes" extend-
ing conspicuously onto the lateral sides of
the abdomen.
There are 34 juvenile and subadult A. b.
sceJestus, with snout-vent lengths between
46 (USNM 193989) and 94 (ASFS
V21699-700). Three juveniles (snout-vent
lengths 46-61) have umbilici still present.
This entire suite of young lizards shows a
remarkable diversity in dorsal pattern.
Even small specimens may be either uni-
color green (usually with a vertical nuchal
white crescent and a white subocular
spot ) , green with three or four yellow body
bands, or there may be many more bands
resulting from the inteiposition of pale
body bands between the primary pale
body bands. One specimen (ASFS \'29296;
snout-vent, length 70, male ) has both pale
body bands and interstitial pale blotching,
whereas another lizard (MCZ 125621;
snout-vent length 86, female) already
shows the adult pattern of several fine pale
crossbands on a green ground. The largest
subadults, however, (ASFS V21699-700;
snout-vent lengths 94, male and female)
are both presently unicolor and show no
indications of the adult body banding.
That a single juvenile may demonstrate a
pattern change is shown by the following
notes on ASFS V28757, a female with a
snout-vent length of 54: "Alive, emerald
green with about foiu' pale yellow cross-
bands on body; dead — seven narrow brown
body bands which are hollowed, and the
dorsal groimd color now pale yellow-
green." The dewlap in young males is or-
ange, in young females from dull yellow
streaked with charcoal to charcoal.
Comparisons. In color and pattern, A.
h. scelestus differs from all other subspe-
cies. No other named population has six
or seven narrow dorsal crossbands; even
multistnippus is much more conspicuously
banded than scelestus and lacks any sort
of dorsal blotching. A. h. scelestus is
known to intergrade with more northern
caeruleolatus and is presumed to meet li-
torisilva. In each case, there is no difficulty
distinguishing the adjacent forms chromat-
ically. A. h. caeruleolatus typically has (in
males) sky-blue blotches along the junc-
tion of the dorsal and ventral colors, and
is prominently crossbanded with three dor-
sal crossbands. A. h. litorisilva is a blotched
lizard, the dorsal colors much more drab
than those of scelestus, tending toward
browns and brownish greens. Perhaps
scelestus most closely resembles multi-
stnippus, but, although both are banded,
the bands in multistruppus are much finer
and much more numerous than the six or
seven pale dorsal crossbars in scelestus.
A. h. scelestus, with modally 2 snout
scales, differs from caeruleolatus, which
has 4 snout scales. In having 7 vertical lor-
eal rows, scelestus differs from caeruleola-
tus, which has 8 rows. In having 5/5 scales
between the interparietal and supraorbital
semicircles, scelestus differs from samanae,
multistruppus, and .mhlimis, all of which
have 4/4. A. h. scelestus has the lowest
138 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
mean of vertical dorsals ( 15.4 ) of all
named subspecies, being most closely ap-
proached by litorisilva (15.9). A. b. sceles-
tus males are larger than those of any other
subspecies ( 180 in scelestus, 158 in litori-
silva, which is second largest) and in fact
this subspecies exceeds all other Hispanio-
lan giant anoles in size, being most closely
approached by male A. r. ricordi, which
reach a snout-vent length of 160.
Discussion. I am uncertain that all
specimens included in scelestus should be
so associated. This is especially true of the
specimen from near Yamasa (ASFS
V28656); this is a juvenile male and its
taxonomic status remains somewhat in
doubt, since it is young. It is also possible
that specimens from Santo Domingo like-
wise are not identical with more eastern
lizards, although the two samples agree
fairly well.
A. b. scelestus and A. b. caeruleolatus in-
tergrade in the region of El Seibo Prov-
ince; I have examined the following mate-
rial from El Seibo which I consider
intergradient: 3.5 mi. (5.6 km) S Sabana
de la Mar (ASFS X7877); 2.1 mi. (3.4 km)
N El Valle (ASFS X7861-62); 3 km N El
Valle (ASFS V3157-58); 10.5 km N Hato
Mayor (ASFS V35329-30). This series
consists of three juveniles and four young
adults (with snout-vent lengths between
112 and 127). The single adult male
(ASFS X7877) was tannish gray in life
with darker brown blotches, a pale green
venter, and an orange dewlap. Two adult
females (ASFS X7861-62) were pale pea-
green with vertical gray bars, the upper
surface of the head grayish tan, venter
green, and dewlap grayish orange. The
lower jaw and throat were green mottled
with darker green. In general this series
seems closer to caeruleolatus than to scel-
estus, but the male lacks sky-blue ventro-
lateral markings. On the other hand, the
vertical gray bars, recorded for the female,
resemble the pattern of scelestus rather
than that of female caeruleolatus. It seems
likely that caeruleolatus and scelestus in-
tergrade in this region.
Remarks. Almost all ASFS scelestus
were secured while the lizards were asleep
at night. Typical situations are lowland
cacaotales and cafetales with their high
canopied shade-trees, along lowland
streams (as at Otra Banda and Yamasa),
and in woods associated with limestone
cliffs (east of Santo Domingo). The long
series from the Rio Cumayasa is from the
high riverine woods along that stream; re-
markably, we secured only juveniles and
subadults at this locality, despite three
nocturnal visits. One juvenile from this lo-
cality was taken on the exposed branch of
an Acacia tree along an open road. Per-
haps the most remarkable place whence A.
b. scelestus has been taken is the locality
northeast of La Romana. This place is a
deep and well-wooded ravine through
which flows a clear stream; however, the
ravine is completely surrounded on all
sides by cane fields, and the ravine woods
are completely isolated at the ravine rim
from other such ecologies, if they even still
exist in this area. A. b. scelestus was ex-
ceptionally abundant in this particular and
very restricted strip of riverine gallery for-
est. Elevations above ground recorded for
sleeping scelestus range from 2 to 20 feet
(0.6 to 6.1 meters), with juveniles usually
sleeping much lower than adults. The alti-
tudinal distribution of A. b. scelestus is in
general low, with recorded elevations from
sea level to 200 feet. It is likely that this
subspecies also occurs in the uplands of
the Cordillera Oriental, but as yet there
are no specimens from areas within that
rather low-lying but mesic and well-for-
ested massif.
The name scelestus is from the Latin for
"unlucky, wretched," in allusion to the dif-
ficulties involved with collecting this sub-
species at the La Romana ravine noted
above.
The transition betu'cen scelestus and //-
torisilva must be very abrupt; the two sub-
species are known from localities separated
by only 28 kilometers. The habitats of the
two subspecies are quite different, with
scelestus inhabiting very mesic situations
HisPANiOLAN Giant Angle • Schwartz 139
and litorisilva xeiic coastal woods. Inter-
estingly, this same eastern region of the
Repiiblica Dominicana is also an area of
abrupt changes in subspecies of Anolis di-
stichus, where the subspecies ifi,ni<iularis
Mertens and properus Schwartz have
ranges which coincide rather closely with
those of scelestus and litorisilva (see
Schwartz, 1968: 275, map). The question
of intcrgradation between scelestus and the
southwestern subspcx-ies next to be named
will be discussed under that taxon.
Perhaps more so than any other subspe-
cies, scelestus seems to show a very spotty
distribution. Two instances are worthy of
mention. There are excellent extensive
coastal forests at Cabo Caucedo south of
the Aeropuerto Internacional de las Ame-
ricas on the southern Dominican coast. Re-
peated diurnal and nocturnal visits to these
splendid woods yielded no A. baleatus, de-
spite what seems to be more than adequate
habitat. A second locality, east of Boca
Chica along the same coast, likewise sup-
ports extensive fine stands of lowland hard-
wood forests, and there also, despite many
diurnal and nocturnal visits, we have never
encountered A. baleatus. It is possible that
these two instances of fairly dry coastal
woods are not suitable for scelestus
(whereas they surely would be for litori-
silva) and that scelestus simply does not
occur there.
Anolis baleatus fraudator
new subspecies
Holotype. USNM 193978, an adult fe-
male, from 4 km W, 6 km N Azua, Azua
Province, Repiiblica Dominicana, one of
two taken by Richard Thomas, on 23 July
1969. Original number V21384.
Paratypes. ASFS V21385, same data as
holotype; ASFS V21433, Barreras, Azua
Province, Repiiblica Dominicana, native
collector, 25 July 1969; ASFS V723, 1.1 mi.
(1.8 km) S San Jose de Ocoa, 1400 feet
(427 meters), Peravia Province, Repiiblica
Dominicana, R. F. Klinikowski, 24 August
1958; ASFS V21203, Sierra Martin Garcia,
about 3000 feet (915 meters), above Bar-
reras, Azua Province, Repiiblica Domini-
cana, R. Thomas, 20 July 1969; ASFS
V31207, Sierra Martin Garcia, above Bar-
reras, between 2000 and 2800 feet (610
and 854 meters), west slope, Mt. Biisi'i,
Barahona Province, Repiiblica Dominicana,
B. R. Sheplan, 15-17 September 1971.'
Definition. A subspecies of A. baleatus
characterized by the combination of mod-
ally 4 scales at level of the second canthal
scales, 6 vertical rows of loreal scales, 2 or
3 scales between the supraorbital semicir-
cles, 4/4 scales between the interparietal
and the supraorbital semicircles, high num-
ber of vertical dorsal scales (17-21; mean
18.8), low number of ventral scales (18-
26; mean 20.7), nuchal scales high, body
crest scales moderate in only adult female,
subocular scales usually separated from
supralabial scales by one row of scales;
dorsum (in female) mottled pale and
darker gray, with three irregular white
crossbands, and blotched with yellow-
green, top of snout and lores straw, labials
dull yellow, and dewlap nearly white with
a yellowish or cream wash.
Distribution. The Sierra Martin Garcia
in Barahona and Azua provinces, and
along the southern slopes of the Cordillera
Central and the Sierra de Ocoa in Azua
and Peravia provinces.
Description of holotype. An adult fe-
male with a snout-vent length of 133 and
tail length of 244; snout scales between
second canthals 4; 6 vertical rows of loreal
scales, 2 scales between supraorbital semi-
1 Since the present manuscript was completed,
a juvenile female ( MCZ 132301) with a snout-
vent length of 57 mm, was secured by E. E. Wil-
liams and J. Roughgarden at a locality south of
La Honna, Peravia Province, on 19 July 1972.
This lizard is to be considered a paratype. It has
3 snout scales at the level of the second canthals,
6 loreal rows, 3 scales between the supraorbital
semicircles, 5/5 scales between the interparietal
and the semicircles, 16 vertical rows of dorsal
scales and 20 rows of ventral scales, and 1 scale
between the suboculars and the supralabials. Both
nuchal and body crest scales are low. As pre-
served, the lizard is dull greenish with indications
of dark dorsal crossbars, and it lacks any pale
dorsal markings.
140 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
circles, 4/3 scales between the interpari-
etal and the supraorbital semicircles, verti-
cal dorsals 17, horizontal dorsals 24,
ventrals 21, subocular scales separated
from supralabial scales by one row of
scales, fourth toe lamellae on phalanges II
and III 33, nuchal crest scales high, body
crest scales moderate; in life, dorsum mot-
tled pale and dark gray, blotched with yel-
low-green and with three irregular white
crossbands, labials dull yellow, top of snout
and lores straw, and dewlap nearly white
with a yellowish or cream wash.
Variation. The only adult is the holo-
type; the remainder of the paratypic series
is composed of juveniles and subadults
with snout-vent lengths between 74 and 96
(three males, two females). Snout scales
at the level of the second canthals range
between 2 and 4; the mode is 4 ( four spec-
imens). The vertical loreal rows vary be-
tween 5 and 7, with a mode of 6 (three
specimens). There are 2 or 3 scales be-
tween the supraorbital semicircles; both
categories have the same frequency. There
are modally 4/4 scales between the inter-
parietal and the supraorbital semicircles; 4
scales are involved in 67 percent of the
combinations; actual counts are 3/4 (2),
4/4 (3), and 5/6 (1). Vertical dorsals
range between 17 and 21 (mean 18.8),
horizontal dorsals between 20 and 24
(21.4), and ventrals between 18 and 26
(20.7). The only adult specimen (a fe-
male) has the nuchal crest scales high and
the dorsal body crest scales moderate. Five
specimens have the suboculars separated
from the supralabials by one row of scales
and one lizard has these scales in contact
( 17 percent ) .
The details of the color and pattern of
the only adult, the female holotype, have
already been given. The juveniles and
subadults show the same general pattern
configuration as does the adult. The small-
est juvenile (snout-vent length 72), a fe-
male topotype, was gray with yellowish
mottling and a pat*"ern of three irregular
crossbands, a faint white scapular stripe,
and black postauricular and postorbital
spots. The dewlap was charcoal with
white scales. The next largest individual
(snout-vent length 74), a male, had the
dorsum pale green with irregular trans-
verse barring; the upper surfaces of the
limbs were pale green and gray-green, and
the tail was banded pale green and gray-
green. The venter was whitish. The dew-
lap was very dark yellowish with an or-
ange wash posteriorly. A slightly larger
male (snout-vent length 85) was pale
green, much marbled and shaded with tan
to gray and with some faint evidence of
transverse crossbands; the chin and throat
were gray-green, and the dewlap pale gray-
ish orange. A female from the Sierra Mar-
tin Garcia (snout-vent length 88) was
green and brown dorsally and without pale
markings; the dewlap was marbled with
charcoal. Finally, the largest subadult
(snout-vent length 76), a male, had the
dewlap dirty yellow with orange streaking.
In the case of fraudator, the very pale ( al-
most white) adult fem.ale dewlap appears
to be preceded ontogenetically by brighter
and more typically A. haleatus hues.
Comparisons. No other subspecies of A.
haleatus approaches the pale colors of
fraudator, nor does any other subspecies
have such a pale dewlap. Although frau-
dator combines the blotching and trans-
verse crossbands in the same fashion as
does scelestus, fraudator is in all ways a
paler lizard. Comparisons in details of
color and pattern with all other subspecies
of A. haleatus are unnecessary. A. h. frau-
dator differs from samaime, scelestus, multi-
struppus, and suhliinis in having 4 rather
than 2 snout scales at the second canthals,
and only fraudator has a mode of 6 verti-
cal rows of loreals (7 or 8 in all other sub-
species). In having 4/4 scales between the
inteiparietal and the supraorbital semicir-
cles, fraudator differs from caeruleolatus,
scelestus, and haleatus, all of which have
5/5. Although fraudator has a high mean
(18.8) of vertical dorsals, in which it is
exceeded only by suhlimis (mean 19.2),
fraudator has the lowest mean (20.7) of
HisPANiOLAN Giant Angle • Schwartz
141
ventrals of all subspecies, being approached
most closely by scelestus (21.1).
Discussion. Apparently A. h. fraudator
is a pale subspecies that is restricted to fa-
vored situations in the xeric regions asso-
ciated with the Llanos de Azua along the
southern slopes of the Cordillera Central
and the Sierra de Ocoa, a southern affiliate
of the former range. The subspecies ap-
parently also occurs in the Sierra Martin
Garcia, an eastern isolate of the Sierra de
Neiba (which, it will be recalled, is else-
where occupied by A. r. ricordi) and sur-
rounded by extreme desert. The specimen
from Barreras, which lies at the foot of the
Sierra Martin Garcia, is interesting in that
it seems a most unlikely locality for any
giant anole; however, I assume that the
specimen, which was native-collected, was
taken either in nearby Cocos groves or on
the lower wooded slopes of the range it-
self. Two specimens from the higher eleva-
tions of the Martin Garcia are from dense
woods, and the specimen from San Jose
de Ocoa was taken from a large tree at the
edge of a pasture. The type locality is
semi-xeric woods with vine tangles and
mango trees in an otherwise cultivated but
xeric region. Probably A. b. fraudutor is
widely distributed in suitable situations
through much of this region, but the lizard
appears to be rare; Buffett and I collected
in semi-mesic riverine woods at a locality
4 km W and 17 km N Azua at an elevation
of about 500 feet ( 153 meters ) , both dur-
ing the day and at night, without seeing
any giant anoles. Natives just south of San
Jose de Ocoa at an elevation of 1400 feet
(427 meters) did not secure specimens for
us in semi-mesic woodlands. Since the al-
titudinal distribution of fraudator extends
from sea level to about 3000 feet (915 me-
ters) in the Sierra Martin Garcia, the ele-
vations of the above-mentioned localities
are within the known altitudinal range of
the subspecies, and indeed our San Jose de
Ocoa locality was quite close to where
Klinikowski secured one of the paratypes.
Remarks. The name fraudator is from
the Latin for "deceiver" in reference to the
resemblances between this sui)species and
A. harahonae. In fact, my decision to re-
gard fraudator as a subspecies of baleatus
rather than harahonae is based more upon
the juveniles than the adults of fraudator;
this is not exclusively due to the fact that
there are more juveniles of fraudator than
adults but rather that the patterns shown
by juvenile fraudator are more typically
those of A. Jjaleatus than of A. harahonae.
A. J), harahomie and A. h. fraudator are
alike in modal numbers of scales at the
level of the second canthals (4), vertical
loreal rows (6), and scales between the in-
terparietal and the supraorbital semicircles
(4/4), and they do not differ strikingly in
means of body scales ( 17.2, 18.8 in vertical
dorsals; 18.2, 21.4 in horizontal dorsals;
22.1, 20.7 in ventrals). In these means,
harahonae is lower in dorsal body counts,
but higher in ventral coimts. The moder-
ate nuchal crest scales of fraudator occur
also in harahonae, but most female hara-
honae have these scales low. No female
harahonae has moderate dorsal body crest
scales as does the female fraudator,
whereas moderate body crest scales occur
in females of most subspecies of A. halea-
ttis ( only female suhhniis lack them ) . Tak-
ing all evidence into consideration, I have
elected to consider fraudator a subspecies
of A. ])aleatus, but its resemblances to A.
harahonae are acknowledged. The distance
separating these two species in this area
is only 20 kilometers (see introduction),
and it is not unlikely that A. harahomie has
been derived from fraudator across the
strait that is now the Valle de Neiba (see
discussion). On the other hand, A. h.
fraudator is removed by some 60 kilometers
from the nearest A. ricordi locality in the
nearby Sierra de Neiba. There is no ques-
tion that fraudator is not correctly associ-
ated nomenclatorially with A. ricordi.
The apparent geographic isolation of
fraudator in relation to other subspecies of
A. haleatus is probably artificial. The near-
est records for other subspecies are: cae-
ruleolatus — 38 kilometers ( San Jose de
Ocoa and La Cumbre); scelestus — 55 kilo-
142 Bulletin Museum of Comparaiive Zoology, Vol. 146, No. 2
meters (San Jose de Ocoa and Yamasa);
and sublimis — 50 kilometers (San Jose de
Ocoa and south of EI Rio). There are
suitable habitats for giant anoles between
caertileolatus, scelestus, and fraudator, but
specimens are lacking. The intervening
high Cordillera Central between the ranges
of sublimis and fraudator probably acts as
a barrier to prevent contact between these
two subspecies.
DISCUSSION
My decision to consider Anolis ricordi
as three species rather than one has some
precedent in the Schwartz and Garrido
(1972) treatment of the Cuban Anolis
equestris, wherein that species was divided
into five species. However, the two situa-
tions, although comparable, are far from
identical. In the A. equestris complex,
there are at least a few incidences of sym-
patry between members of the species-
complex which give clues to the facts of
the situation; there are sti'ong differences
in size of dorsal scales; there are some
strong differences between details of pat-
tern and coloration of the axillary stripe
and the dewlap which likewise suggest
that we are there dealing with more than
one species. But on the other hand, the
Hispaniolan giant anoles show absolute
differences in the nuchal and body crest
scales and differences in the pattern of the
body itself, as well as modal differences in
scutellar details. In addition, there are no
cases as yet known in Hispaniola of sym-
patry between the three entities that I re-
gard as full species. The gaps between
them are narrow, however, and I feel
strongly that it is merely a matter of get-
ting into the intermediate areas and, once
there, being fortunate enough to encounter
giant anoles.
It should be obvious from my systematic
treatment that I am convinced that we are
dealing in Hispaniola with three distinct
species — ricordi, barahonae, and baleatus.
Surely the differences between ricordi and
baleatus are such that, when taken in sum,
one has no doubts that he is involved with
two very different animals. The differences
here are much greater, for instance, than
between Anolis distichus Cope and Anolis
hrevirostris Bocourt, two species that were
long confused and that resemble each
other moiphologically to a very great de-
gree. Yet once one learns what the char-
acters are for separating them, he experi-
ences little difficulty in dealing with both
populations or individuals, either alio- or
sympatric, of these two species. The dif-
ferences in life, as far as pattern and color
are concerned, are not particularly subtle,
and the details of scutellation are not di-
chotomous, but the modal differences are
so well correlated with the pigmental and
pattern traits that we now recognize these
two species with assurance.
An even more obvious parallel is Anolis
carolinensis Voigt and Anolis allisoni Bar-
bour in Cuba. These two species of green
anoles, long confused as A. porcatus Gray
(or A. c. porcatus), were shown by Ruibal
and Williams (1961) to be a sibling pair,
fairly allopatric but both widely distrib-
uted throughout much of Cuba, and to
differ structurally by the condition of the
postauricular area. The presence (allisoni)
or absence (carolinensis) of a deep and
elongate postauricular groove in these two
species is correlated with very striking dif-
ferences in adult pattern and coloration
and other details of scutellation.
The same situation, that of two species
masquerading under a single name, can
also be demonstrated in Anolis alutaceus
Cope and Anolis clivicola Barbour and
Shreve (Schwartz and Garrido, 1971), and
the two species recently confused under
Anolis spectrum Peters; both these situa-
tions pertain to Cuban species. Sr. Gar-
rido also advises me that he has much
evidence to indicate that Anolis cyanopleu-
rus cupeyalensis Peters is in fact a sym-
patric sibling, rather than a subspecies, of
A. cyanopleurus Cope.
I could cite other examples in Antillean
iguanids (Leiocephalus) and anguids
(Diploglossus) which demonstrate quite
clearly the above trend. As more material
HisPAXioLAN Giant Angle • Schwartz
143
from more diverse localities becomes avail-
able, and as this material is subjected to
re-evaluation with differing and more mod-
ern philosophies, our impressions of rela-
tionships among Antillean anolines have
been modified or changed. A major factor
in such revisions has invariably been a
great quantity of new material from areas
that had previously been unsampled, cou-
pled with pigmental, ecological, and etho-
logical data from the living specimens. A
second general line of evidence, equal to
or possibly suipassing morphological and
distributional data in importance, is karyo-
typic and electrophoretic information. One
or both of these areas of investigation are
increasing our knowledge of the complex-
ities within such a genus as Anolis. When
these two areas of research — morphologi-
cal and biochemical — can be brought to
bear simultaneously upon a single species
or species complex, the results may be even
more meaningful than either is alone. As
yet this has not been done in any of the
Antillean giant anoles, so that my conclu-
sions, based upon morphology and distri-
bution, remain to be verified by other evi-
dence. Yet I feel as secure as any
systematist can be when he is dealing with
data that are incomplete.
As pointed out in the introduction to the
present paper, the taxa ricordi, haleatus,
harahonae, and leberi are, on inspection,
unequivocally distinct. But the degree or
level of differentiation of these four taxa
seems to be two-fold. On one hand (ri-
cordi and haleatus), the two populations
are easily separable on the basis of a struc-
tural feature (the nuchal crest scales), a
character that is strongly correlated with
obvious pigmental and pattern traits. On
the other hand, the differences between
harahonae or leheri and ricordi are pri-
marily ones of pigmentation and pattern,
with morphological differences much less
trenchant than between ricordi and ha-
leatus. At the outset such a dichotomy sug-
gests that it might be more proper to con-
sider "A. ricordi" as a complex of full
species than as one species with four (or
more) subspecies. Appar(Mitly Williams
and Rand (1969) had the same inclina-
tions, since they indicated that the differ-
ences between some of the then-named
populations of A. ricordi were such as to
suggest that there might be more than one
species involved.
Once the above assumption has been
made — namely, that A. ricordi is composed
of more than one species — then the prob-
lem first becomes one of differentiating and
delimiting the component species. There
is no difficulty here in separating A. ricordi
and A. haleatus on the basis of crest scales.
None of the populations of A. haleatus has
the moderate (rarely) to low (usually)
nuchal crest scales of A. r. ricordi. In ad-
dition, the narrow geographical gaps that
exist between A. ricordi and A. haleatus
also suggest that these two taxa may be
either allopatric or may meet and occur
sympatrically without intergradation.
The status of the Tiburon populations
that I associate nomenclatorially with A.
ricordi and that of A. harahonae as a dis-
tinct species are less clear than the ricordi-
haleatus relationship. First, the named
populations leheri, viculus, and suhsolamis
have in common a suite of pattern and
color features that ally them more closely
to each other than to A. r. ricordi. The
only evidence for this relationship is the
occurrence of presumed viculus X ricordi
intergrades in the Miragoane-Paillant re-
gion. Were it not for these specimens, I
would be strongly tempted to consider the
three Tiburon taxa as comprising a sepa-
rate species. Any interpretation of the re-
lationships of the Tiburon taxa suffers
from paucity of material from a variety of
localities.
The situation with A. harahomie is in
some ways puzzling. Although there is no
question that it is distinct from A. ricordi,
its relationships to A. haleatus are much
less certain. This uncertainty is caused by
A. h. fraudator, that population assigned to
A. haleatus which is closest geographically
to A. harahonae. It is particularly unfortu-
nate that fraudator is known from only
144 Bulletin Museum of Coinparative Zoologij, Yo]. 146, No. 2
one adult and several juvenile and sub-
adult specimens, since adult males ( primar-
ily) would be most instructive in compar-
ing frattdator with barahonae. On the
other hand, the closeness of fraiidator and
barahonae in characteristics may be rather
a reflection of the ancestry of A. bara-
honae— namely, that it is a south island
(sensu Williams, 1961) invader from the
north, and that the parent population has
been fraudator rather than any other sub-
species of A. baleatus or A. ricordi from the
west.
It might be more proper either to con-
sider A. barahonae as conspecific with A.
baleatus (the two taxa linked through
fraudator) , or to consider fraudator a sub-
species of A. barahonae; either interpreta-
tion has merit. The course that I have
taken seems satisfactory at the moment but
surely is subject to reinterpretation with
the acquisition of more material from this
critical geographic area.
The history of the Hispaniolan giant
anoles appears to be correlated with the
two palaeo-islands that have been fused at
the level of the Cul de Sac-Valle de Neiba
plain with lowering Pleistocene sea levels.
I suggest the following history for the com-
plex; the reader should keep in mind that
such a history is based upon taxonomic
premises that are inductive, and the cau-
tions and uncertainties that I expressed
above have special application here.
Distributional evidence suggests that the
giant Hispaniolan anole stock was origi-
nally restricted to the north island (north
of the Cul de Sac-Valle de Neiba plain).
In this region, two distinctive species arose,
ricordi in the west and baleatus in the east.
There apparently has been local differen-
tiation on the north island at a subspecific
level in both these species, but that in ri-
cordi remains unanalyzed because of too
few specimens. On the other hand, differ-
entiation in A. baleatus is now fairly well
known and documented. This species oc-
curs east of the Cordillera Central and on
the southern slopes of that range and in
the Sierra Martin Garcia. There have been
local population differentiations in response
to the various ecologies within the area oc-
cupied, with two major integumental
trends (coloration and pattern) and details
of scutellation of the head and body (al-
though the latter is not so clear as the for-
mer ) .
There seem to have been two subse-
quent invasions of the south island. To the
west, a (presumably) early invasion of the
A. ricordi stock crossed what is now the
Cul de Sac Plain into the Port-au-Prince
area. It is pertinent that many north island
species have made this same crossing and
have extended their ranges but little fur-
ther. These species with more restricted
ranges have been handicapped either by
competition with already established spe-
cies, improper ecological situations, or rela-
tively recent arrival. A. ricordi seems to
have been an early arrival, without local
competitors, and with abundant proper
ecology (mesic forests). The species has
thus expanded its range after the original
crossing to cover the entire Tiburon Penin-
sula, having somewhere succeeded in cross-
ing the mountainous spine of the Massif de
la Hotte-Massif de la Selle. Local differ-
entiation along the Tiburon in response to
lack of genetic contact across the interior
mountains has also taken place. Further
speculations on details of the history of A.
ricordi on the Tiburon Peninsula are point-
less, since the specimens upon which any
generalisations may be made do not as yet
exist in collections.
A second invasion to the east occurred
presumably at a later date, after the estab-
lishment of A. ricordi on the Tiburon Pen-
insula. This latter invasion resulted in the
differentiation of A. barahonae (from a
fraudator or pre-fraudator stock on the
southern portion of the north island ) in the
Sierra de Baoruco and its subsequent ex-
pansion onto the southern portion of the
Peninsula de Barahona and east along the
southern slopes of the Sierra de Baoruco.
With the previous establishment of A. ri-
cordi to the west (as at Thiotte), the
western movement of A. barahonae was
HisPANiOLAN Giant Angle • Schwartz
145
halted by the presence of the related spe-
cies. I have no doubt that both A. ricordi
and A. barahonae will be found to be
closely allopatric or synipatric in extreme
southeastern Haiti between the Dominico-
Haitian border and Saltrou, and also that
these two species meet and interact along
the northern slopes of the Sierra de Bao-
ruco and the Morne des Enfants Perdus.
One other distributional detail requires
comment. The occurrence of A. r. ricordi
in the main mass of the Sierra de Neiba on
the northern side of the Valle de Neiba
and of A. h. fraudator in the Sierra Martin
Garcia, an extreme eastern isolate of the
Sierra de Neiba, has already been noted.
The Martin Garcia seems to have been
long isolated from not only the Sierra de
Neiba but also from all other Hispaniolan
mountain masses; it is ideally a montane
island in a sea of desert. It seems likely
that this range was unoccupied by giant
anoles of either species (A. ricordi or A.
baleatus), despite the fact that the range
forms a portion of the Neiba uplift. Inva-
sion of the Martin Garcia was possible
from either the northwest (ricordi) or the
northeast (baleatus). Of the two species,
A. baleatus was the more vagile and
reached the Sierra Martin Garcia from the
relatively more mesic southern slopes of
the Cordillera Central before A. ricordi
reached it across the deserts and xeric hills
between the Sierra de Neiba and the Mar-
tin Garcia. This upland population in
turn was responsible for the invasion of the
Sierra de Baoruco across the better for-
ested and more mesic eastern end of the
Valle de Neiba.
Wetmore and Swales (1931: 235) re-
ported the finding of recent Anolis ricordi
skeletal material in Barn Owl (Tyto alba)
pellets from L'Acul, Dept. du Sud, Haiti,
on the Tiburon Peninsula, and Hecht
(1951: 23) noted the abimdant remains of
the species from deposits in "Deep Cave,"
near St. Michel de I'Atalaye, Dept. de I'Ar-
tibonite, Haiti. Etheridge (1965: 101) re-
ported A. ricordi remains from recent owl
pellets near the mouth of a cave near Boca
de Yuma, La Altagracia Province, Repu-
blica Dominicana. Etheridge (op. cit.: 87-
88) also noted pre-Columbian giant anole
remains from a cave at Cerro de San Fran-
cisco near Pedro Santana, La Estrelleta
Province, Repiiblica Dominicana. From the
suite of about 80 cranial elements and
eight pelves, Etheridge extrapolated that
the maximally sized individuals in the cave
deposits had a snout-vent length of 190-
192 mm, some 30 mm larger than any liv-
ing A. ricordi recorded ( 159 mm, fide Eth-
eridge, op. cit.: 88). The maximally sized
Hispaniolan giant anole recorded in the
present paper reaches a length of 180 mm
(male A. b. scelestus from Otra Banda, La
Altagracia Province, Republica Domini-
cana). The difference between this mod-
ern living lizard and the maximally sized
pre-Columbian lizards is not so great as
Etheridge's data suggest. Intriguingly, the
Cerro de San Francisco area lies within the
known range of A. r. ricordi, and the larg-
est specimens of this subspecies ( male with
a snout-vent length of 160 mm, female 151
mm) are from the southern slopes of the
Cordillera Central, veiy close to the Cerro
de San Francisco area. Although there
seems to have been some change in maxi-
mum size in Hispaniolan giant anoles with
the passage of time, these changes have
not been of the magnitude that previous
data suggested.
LITERATURE CITED
Cochran, D. M. 1941. The herpetology of
Hispaniola. Bull. U.S. Natl. Mus., 177: 398,
120 figs., 12 pis.
Cope, E. D. 1864. Contributions to the herpe-
tology of tropical America. Proc. Acad. Nat.
Sci. Philadelphia, pp. 166-181.
DUMERIL, A. M. C, AND G. BiBRO.N. 1837.
Erpetologie gcnerale ou histoire naturelle
complete des reptiles, vol. 4. 571 pp., 14 pis.
Etheridge, R. E. 1965. Fossil lizards from the
Dominican Republic. Quart. Jour. Florida
Acad. Sci., 28(1): 83-195, 3 figs.
Garrido, O. H., AND A. Schwartz. 1968. Cu-
ban lizards of the genus Chamaeleolis. Quart.
Jour. Florida Acad. Sci., 30(3): 197-220, 2
figs.
Hecht, M. K. 1951. Fossil lizards of the West
146 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2
Indian genus Aristelliger ( Gekkonidae ) .
Amer. Mus. Novitates, No. 1538: 1-33, 8 figs.
Maerz, a., and M. R. Paul. 1950. A Diction-
ary of Color. New York: McGraw-Hill Book
Co., pp. vii + 1-23, 137-108, 56 pis.
Mertens, R. 1939. Herpetologische Ergebnisse
einer Reise nach der Insel Hispaniola, West-
indien. Abh. senckenberg. naturf. Ges., 449:
1-84, 10 pis.
Rand, A. S., and E. E. Williams. 1969. The
anoles of La Palma; aspects of their ecolog-
ical relationships. Breviora, Mus. Comp.
Zool., No. 327: 1-18, 1 fig.
RuiBAL, R., AND E. E. Williams. 1961. Two
sympatric Cuban anoles of the carolinensis
group. Bull. Mus. Comp. Zool., 125(7):
183-208, 11 figs.
Schmidt, K. P. 1921. Notes on the herpetology
of Santo Domingo. Bull. Amer. Mus. Nat.
Hist., 44(11): 7-20, 12 figs.
Schwartz, A. 1964. Anolis equestris in Ori-
ente Province, Cuba. Bull. Mus. Comp.
Zool., 131(12): 407-428, 7 figs.
. 1968. Geographic variation in Anolis
distichus Cope ( Lacertilia, Iguanidae ) in the
Bahama Islands and Hispaniola. Bull. Mus.
Comp. Zool., 137(2): 255-309, 4 figs., 2 pis.
, and O. H. Garrido. 1971. The status
of Anolis alutaceus clivicolus Barbour and
Shreve. Caribbean Jour. Sci., 11(1-2): 11-
15.
. 1972. The lizards of the
AND
Anolis equestris complex in Cuba. Stud.
Fauna Curasao and Caribbean Is., 39(134):
1-86, 8 figs.
Thomas, R. 1971. A new species of Diploglos-
sus (Sauria: Anguidae) from Hispaniola.
Occ. Papers Mus. Zool., Louisiana State Univ.,
40: 1-9, 4 figs.
Wetmore, A., and B. H. Swales. 1931. The
birds of Haiti and the Dominican Republic.
Bull. U.S. Natl. Mus., 155: 1-483, 2 figs., 26
pis.
Williams, E. E. 1961. The evolution and rela-
tionships of the Anolis semilineatus group.
Breviora, Mus. Comp. Zool., No. 138: 1-8, 1
pi.
. 1962. Notes on Hispaniolan herpetol-
ogy. 6. The giant anoles. Breviora, Mus.
Comp. Zool., No. 155: 1-15, 1 fig.
. 1965. Hispaniolan giant anoles (Sau-
ria, Iguanidae ) : new data and a new subspe-
cies. Breviora, Mus. Comp. Zool., No. 232:
1-7, 2 figs.
, and a. S. Rand. 1969. Anolis insolitus, a
new dwarf anole of zoogeographic importance
from the mountains of the Dominican Repub-
lic. Breviora, Mus. Comp. Zool., No. 326:
1-21, 6 figs.
us ISSN 0027-4100
BulLetln OF TH
seum
Comparative
Zoology
A Revision of the Cardinalfish Genus
Epigonus (Perciformes, Apogonidae)^
with Descriptions of Two New Species
GARRY F. MAYER
HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSETTS, U.S.A.
VOLUME 146, NUMBER 3
19 SEPTEMBER 1974
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OccAsioNAL Papers on Mollusks, 1945-
SPECIAL PUBLICATIONS.
1. Whittington, H. B., and E. D. I. Rolfe (eds.), 1963. Phylogeny and
Evolution of Crustacea. 192 pp.
2. Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredini-
dae (Mollusca: Bivalvia). 265 pp.
3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echinoderms.
284 pp.
4. Eaton, R. J. E., 1974. A Flora of Concord. 211 pp.
Other Publications.
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Reprint.
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Insects.
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Mammalian Hibernation.
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Proceedings of the New England Zoological Club 1899-1948. (Complete
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Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massa-
chusetts, 02138, U.S.A.
© The President and Fellows of Harvard College 1974.
t
A REVISION OF THE CARDINALFISH GENUS EPIGONUS
(PERCIFORMES, APOGONIDAE), WITH DESCRIPTIONS
OF TWO NEW SPECIES^
GARRY F. MAYERS
TABLE OF CONTENTS resim-ected and redescribed on the basis of new
. ,^Y material, and Htjnnodus atherinoides Gilbert and
:^ j*^ " , ,« H. megalops Smith and Radcliffe are synonymized
Introduction 14^ ^^^^^j^ ^ occidentaUs Goode and Bean.
Metnods Species descriptions include discussions of dis-
Systematics """r " ici tribution, geographic variation, ontogenetic
Genus Epigonus Rafmesque 151 ^^^^^^^ a„j taxonomic problems. An in%'estiga-
Diagnosis 151 ^^^^ ^^ ^j^^ ^^p^^ ^^j ^ lenimen (Whitley) reveals
Description 151 ^^^^^ ^^ holotype and paratypes are not conspe-
Key to the Species of Epigonus 152 ^.jjj^ Instead, the paratypes are members of E.
Epigonus telescopus 152 denticulatus Dieuzeide. A key to the species of
Epigomis macrops 159 Epigonus is provided at the beginning of the
E))igonus ))andionis 163 paper.
Epigonus fragilis 169
Epigonus occidentaUs 170 INTRODUCTION
Epigonus denticulatus 1^5
Epigonus oligolepis 179 Selected species of Epigomis have been
Epigonus tretvavasae 183 j^^^^^^^^^ 1^ ^^^.^^^ f^^ at least two hundred
Epigonus pectimfcr loo J , , onr, r>^\ i j
Epigonus wbustus 1S9 ti^ty Y^ars. Vaillant (1888: 25) remarked
Epigonus lenimen 193 that E. telescopus was recognized in ancient
Epigonus crassicaudus 197 times, and Risso (1810: 303) reported that
Species Incertae Hedis 199 j-j-jj^ gpecies was prized for its firm, deli-
Aek„S^!Sr°!!!' ::;:::::::::::::::::::::::::: loE! cious-tasti„g „,«,, although it was rardy
Literature Cited 200 taken. The presence of common names tor
Appendix 203 E. telescopus in vocabularies of western
Mediterranean and North Atlantic fishing
Abstr.\ct. a study of the deep-sea Apogonidae communities ( Doderlein, 1889 ) provides
results in a revision of the genus Epigonus additional evidence of man's long-term
Rafinesque. Twelve species are recognized in- awareness of the species. £. telescopus is
eluding two new forms — E. oligolepis and £...,, • n i i • . i i i. „r
pectirUfer. E. fragilis (Jordan and Jordan) is still occasionally sold in the markets of
southwestern Europe.
' This paper is based on a portion of a thesis Two other species of EpigOnus are cap-
presented to Harvard University in partial ful- tured by commercial fishermen. E. denti-
fillment of the requirements for the Ph.D. in culcitUS is edible ( Dieuzeide et al., 1953:
^i?'"g>- ^ r -1^) and is taken in the Mediterranean.
- Department of Marine Science, University of y-r .., .i ,i . ,■ ^^t■^^■ >., f^^
c u \ri J c^ u . I 171 • 1 oo-m 1 Uutil recently this form was mistaken tor
South Florida, St. Petersburg, Florida 33/01 and ^■'^_, , _, , ,
Museum of Comparative Zoology, Harvard Uni- the yOUng of E. teUsCOpUS. E. crassicaudus
\ersity, Cambridge, Massachusetts 02138. is caught by Chilean fishermen. Like E.
Bull. Mus. Comp. Zool., 146(3): 147-203, September, 1974 147
148 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
telescopus, it is not taken in sufficient
numbers to support a separate fishery but is
captured by fishermen trawhng for more
abundant deepwater organisms.
Although African Epigonus are not pres-
ently exploited, tropical eastern Atlantic
stocks may represent future sources of pro-
tein for mankind. Surveys sponsored by
the Organisation of African Unity and the
U.S. Agency for International Development
revealed these fishes are "of possible po-
tential importance ( not necessarily by pres-
ent marketing standards) [Williams, 1968:
79]." The same may be true for Caribbean
and Gulf of Mexico Epigonus; however,
complete data have not been compiled for
the latter areas.
A major hindrance to the evaluation of
deep-sea cardinalfish stocks has been taxo-
nomic confusion. The systematic history of
Epigonus began in 1810 with Risso's de-
scription of Pomatomus telescopus and
Rafinesque's account of its synonym Epi-
gonus niacrophthahnus. During the fol-
lowing seventy-one years, work on the
genus was primarily limited to re-descrip-
tions of E. telescopus and discussions of its
biology (e.g., Cuvier, 1828; Valenciennes,
1830; Capello, 1868; Moreau, 1881).
The surge in oceanographic exploration
during the last quarter of the nineteenth
and beginning of the twentieth centuries
rapidly increased the number of nominal
Epigonus AAke species. Among the forms
described between 1881 and 1920 were
Apogon pandionis Goode and Bean, 1881;
E. occidentalis Goode and Bean, 1896;
Hynnodus atherinoides Gilbert, 1905; Oxij-
odon macrops Brauer, 1906; and Hynnodus
me galops Smith and Radcliffe, 1912. In
the following decade, three new species
and two new genera appeared in the liter-
ature.
Much of the confusion associated with
the taxonomy of Epigonus stems from ma-
terial described prior to 1930. Early taxa
were based on small samples. Because
many nations participated in oceanographic
research, specimens were deposited in
scattered institutions and descriptions ap-
peared in diverse publications. Conse-
quently, it was difficult for workers to
obtain either comparative material or a
broad overview of the group's systematics.
These shortcomings were aggravated by
inaccurate, under-illustrated descriptions
based on ill-considered characters. It was
common, for example, to use dentition
patterns to define generic boundaries, yet
tooth arrangements are difficult to observe,
easily damaged, and subject to ontogenetic
and geographic variation. As a result, an
inordinately large number of Epigonu.s-\ike
forms was recognized by the end of the
1920's.
Although generic taxonomy was stream-
lined by Fowler and Bean in 1930 and
Matsubara in 1936, species-level taxonomy
became increasingly complex. New forms
were described in 1935, 1950, 1954, and
1959. In addition, misidentifications of
Epigonus were published in several widely
circulated works on regional faunas (e.g..
Smith, 1949b and 1961; Gosline and Brock,
1960).
The aim of the present study is to clarify
the species-level systematics of the Epi-
gonus-\ike fishes. Data from traditional
characters are evaluated and augmented
by information from characters not pre-
viously examined for this group. A special
effort is made to discuss features such as
dentition patterns that caused taxonomic
confusion in the past. The ecology, func-
tional anatomy, zoogeography, and evolu-
tion of Epigonus will be discussed in future
works on the genus.
METHODS
Measurements were made to the nearest
tenth of a millimeter by tlie use of Helios
needlepoint dial calipers; characters larger
than 190 mm were measured with a meter
rule or GPM Anthropometer. Measure-
ments routinely taken include:
Standard length (SL) — from tip of snout
to base of caudal fin.
EriaoNus Systematics • Mayer 149
Head length (HL) — from tip of snout to
tip of opercular spine.
Body depth — between dorsal and v(Mitral
surfaces of body at level of peKic fin
base.
Head height — from quadratomandibular
joint vertically to bony rim above eye.
Eye diameter — between anterior and pos-
terior margins of orbit as defined by
first and sixth suborbitals.
Snout length — from tip of snout to an-
terior margin of orbit.
Interorbital width — shortest distance be-
tween bony rims above eyes.
Maxillary length — from tip of snout to
posterior margin of maxilla.
Lower jaw length — from tip of mandible
to quadratomandibular joint.
Caudal peduncle depth — shortest dis-
tance loetween dorsal and ventral sur-
faces of caudal peduncle.
Caudal peduncle length — from posterior-
most anal fin ray to caudal fin base.
First spine length (first spine of first
dorsal fin, D,I; first spine of second
dorsal fin, DJ; second spine of anal
fin, AH; pelvic fin spine, PJ) — from
base to tip of spine along anterior edge.
Counts were made under a dissecting
microscope with the use of dissecting
needles or insect pins. A Fibre-Lite High
Intensity Illuminator proved invaluable for
examinations of oral, branchial, and visceral
structures. Gill raker and branchiostegal
counts were made on the left side of speci-
mens; remaining counts and measurements
were made on the right side whenever
possible. Counts made include: fin spines
(indicated by Roman numerals), fin rays
(indicated by Arabic numerals), branchi-
ostegal rays, rakers on first gill arch, lateral
line scales, pyloric caeca, vertebrae (pre-
caudal + caudal, including hypural fan),
pleural and epipleural ribs, and basal
ptervgiophores between neural spines 9
and 10.
Osteological data were obtained from
radiographs taken at the Woods Hole
Oceanographic Institution, the Museum of
Comparative Zoology, and the Harvard
University School of Public Health. Holo-
types of Oxyodon iiiacrops and Scepterias
Icninwn were radiographed at the Zoolo-
gisches Museum der Humboldt Universitat
and Australian Museum, respectively. More
comprehensive osteological studies were
based on cleared and stained specimens
prepared by trypsin digestion (Taylor,
1967). Osteological terminology follows
that presented by Gosline ( 1961 ) and
Mead and Bradbury (1963). Suborbital
bones are numbered from 1 to 8 beginning
with the rostralmost element (lacrimal).
Statistical data were analyzed with the
use of the Harvard Computation Labo-
ratory's IBM 360/65 digital computer.
Standard techniques described by Mayr
(1969: 189-193) and Simpson et al. (1960:
65-68, 83-88) were employed for analyzing
meristic data. Morphometric characters
were examined with the aid of regression
techniques specified by Simpson et al.
(1960: 215-233, 238) and Bailey (1959:
91-99).
Before undertaking regression analyses,
morphometric data were plotted against
SL. Graphs were drawn according to a
BMD 05D plotting routine (Dixon, 1967:
71 ) and served as visual tests for linearity
of scatter. Only characters exhibiting linear
scatters were analyzed by regression tech-
niques. As a second precaution against
nonlinearity, subadult specimens ( < 40
mm SL) were excluded from statistical
samples.
Data from several morphometric char-
acters are presented both as ratios (i.e.,
percent of SL or HL) and as regression
parameters. The former are intended only
as identification aids. As Royce (1957: 17)
points out, heterogenic growth makes the
use of ratios in fish taxonomy inefficient
and may lead to erroneous conclusions.
Collection and institution names are ab-
breviated as follows in this paper:
ABE —Collection of Dr. T. Abe,
Tokyo
AM — Australian Museum, Sydney
150 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
BMNH — British Museum (Natural
History), London
BPBM — Bernice P. Bishop Museum,
Honohihi
CM — Carnegie Museum; collections
presently housed in FMNH,
Chicago
DM — Dominion Museum, Welling-
ton
EBM — Estacion de Biologia Marina,
Universidad de Chile, Viiia
del Mar
FMNH —Field Museum of Natural
History, Chicago
IRSN — Institut Royal des Sciences
Naturelles de Belgique, Brus-
sels
ISH — Institut fiir Seefischerei, Ham-
burg
LACM — Los Angeles County Museum
of Natural History, Los Ange-
les
MCZ — Museum of Camparative Zo-
ology, Harvard University,
Cambridge
MNHN —Museum National d'Histoii-e
Naturelle, Paris
MZF — Museo Zoologico di Firenze,
Florence
RUSI — J.L.B. Smith Institute of Ich-
thyology, Rhodes University,
Grahamstown
SAM — South African Museum, Cape
Town
SMF — Natur-Museum Senckenberg,
Frankfurt am Main
SU — Stanford University; collec-
tions presently housed in the
California Academy of Sci-
ences, San Francisco
TABL — Tropical Atlantic Biological
Laboratory, Miami
UMML — Rosenstiel School of Marine
and Atmospheric Science,
University of Miami, Miami
USNM — National Museum of Natural
History, Washington, D.C.
UZM — Universitetets Zoologiske Mu-
seum, Copenhagen
ZMB — Zoologisches Museum der
Humboldt Universitiit, Berlin
Descriptions are based on material listed
by Mayer (1972: Appendix II). Additional
data were obtained from examinations of
the seventeen specimens listed below. All
seventeen fishes were radiographed.
E. robustus: ISH 1132/66, 3 specimens,
121.1-142.5 mm, WALTHER HER-
WIG Sta. 237/66, 36°00S, 52°58'W,
800 m. ISH 189/71, 9 specimens,
147.0-198.0 mm, WALTHER HER-
WIG Sta. 121/71, 37°44S, 54°43'W,
800 m. ISH 269/71, 1 specimen, 147.5
mm, WALTHER HERWIG Sta. 340/
71, 38°50'S, 54°25'W, 1000 m. ISH
430/71, 1 specimen, 124.1 mm,
WALTHER HERWIG Sta. 348/71,
38°20'S, 54°33 W, 997-1040 m.
E. fmgilis: LACM 32668-6, 1 specimen,
72.5 mm, 2 mi. off Haleiwa, Oahu,
Hawaii, 65 fms. SU 32262, 2 speci-
mens, 90.0-93.9 mm, Honolulu, Hawaii.
Distributions were determined from
material examined and from published
accounts. Because of the confusion in
Epigomis taxonomy, published data were
used only if species identifications could
be verified from included descriptions,
illustrations, etc. Data from specimens of
doubtful identity were not considered. A
complete list of station data taken from the
literature is provided by Mayer (1972:
Appendix II ) .
No attempt has been made to provide
exhaustive synonymies for Epigonus spe-
cies. References are cited only if they (1)
are taxonomically or zoogeographically im-
portant; (2) provide outstanding descrip-
tions, illustrations, or synonymies; or (3)
represent verifiable misidentifications. Non-
taxonomic accounts have been omitted, as
liave references to cruise summaries and
faunal lists.
SYSTEMATICS
Statistical data are presented in tables
accompanying species descriptions; meristic
Epigonus Systematics • Mayer 151
characters witli low variability are reported
ill the text as value, followed in parentheses
by number of specimens exiiibiting that
v^alue. Meristic and mensural data from
holotypes of new species are presented in
the Appendix.
Genus Epigonus Rafinesque, 1810
Epifiomis Rafinesque, 1810: 64. (Type .species:
Epigonus macrophthahnus Rafinesque, 1810
by in()n()t>p\'. A synonym of Pomatomus
telescopus Risso, 1810.)
Tt'h'scops Bleeker, 1876: 261. (Type .species:
Poiuatonniti tclescopiuiu [sic!] Risso, 1810 by
original designation. Pomatomus deemed in-
applicable. )
Pomatomichthijs Ciglioli, 1880: 20. (Type species:
Pomatomiclithys constanciae Giglioli, 1880 by
monotypy. A synonym of Pomatomus teles-
copus Risso, 1810.)
Hynnodiis Gilbert, 1905: 217. (Type species:
Hijnnodus athcrinoides Gilbert, 1905 by mono-
typy. A synonym of Epigonus occidentaJis
Goode and Bean, 1896.)
Oxyodon Brauer, 1906: 287. (Type species:
Oxyodou Diacwps Brauer, 1906 by monotypy.)
Xystramia Jordan, 1917: 46. (Type species:
Glossamia pandionis Goode and Bean, 1881
by original designation. Glossamia deemed
inapplicable. )
Scepterias Jordan and Jordan, 1922: 44. (Type
species: Scepterias fragilis Jordan and Jordan,
1922 by monotypy.)
Paraliynnodus Barnard, 1927: 525. (Type species:
Parahynnodus robustus Barnard, 1927 by mono-
typy- )
Diapiosis. Epip^onus is distinguished
from other lower perciform genera by a
mosaic of characters including 8 suborbital
bones, all lacking subocular shelves; large,
thin-walled swimbladders with postero-
dorsal ovals; VII or VIII first dorsal fin
spines; 1,9 or 1,10 second dorsal fin ele-
ments; 11,9 anal fin elements; 15-23
pectoral fin rays; and 17-35 gill rakers. No
member of the genus exhibits fang-like
conical teeth, such as are found in Cheilo-
dipterus, or anteriorly projecting teeth,
such as are found in Rosenblattia.
Description. Body elongate, fusiform;
dorsal and ventral profiles slightly convex,
similar. Mouth oblicfue, terminal; upper
jaw protrusile; maxilla excluded from gape.
sheathed l)y lacrimal anteriorly, free pos-
teriorly; supramaxilla absent. Eye large,
round or oval. Nostrils paired, rounded or
slit-like, two on each side of head.
Premaxillae, mandibles, vomer, and pahi-
tines edentulous or bearing conical teeth;
tongue and endopterygoids rarely dc>ntiger-
ous; ectopterygoids edentulous. Gill rakers
moderate to long, 17-35; branchiostegal
rays 7 (3 + 4); pseudobranchiae present.
Opercular .spine either weak, flattened
and poorly ossified, or pungent and bony;
spine ventral to one or more horny or mem-
branous spinelets. Preopercle with double
edge; angle frequently produced.
Dorsal fins VII-1,9, VII-1,10, or VIII-
1,10, separated by distinct interdorsal
space; rudimentary subcutaneous eighth
spine present in seven-spined forms. Anal
fin 11,9; pectoral fins 15-23; peKic fins 1,5:
caudal fin forked, 9 + S principal rays,
upper- and lowermost rays unbranched.
Scales large, deciduous, ctenoid. Lateral
line complete, extending parallel to dorsal
profile on dorsolateral surface of trunk,
descending to midline on posterior portion
of caudal peduncle, continuing on tail;
lateral line scales 33-51; canal simple,
broadening into deltoid or Y-shaped tube
at rear edge of scale. Scale pockets cover-
ing most of body including occiput, soft
dorsal, anal, and caudal fins; scales absent
from snout; no axillary scale at base of
PJ spine.
Suborbitals 8, all lacking subocular
shelves. Vertebrae 25; basapophyses on
vertebrae 3 or 4. Predorsals 3, first and
second interdigitating between neural
spines 2 and 3, third located behind neural
spine 3. Caudal skeleton with 2 autogenous
haemal .spines, 6 hypurals (hypural 1 =
parhypural sensu Monod, 1968), 3 (>purals,
2 (rarely 1) pairs of uroneurals. Actinosts
4, 3VL' borne by scapula.
Swimbladder large, thin-walled, lacking
anterior or posterior projections to cranium
and neural arches; diaphragm absent; oval
posterodorsal; retia mirabilia well devel-
oped. Stomach U- or Y-shaped; pyloric
152 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
caeca .5-34, may be modified into lu-
minescent organs; intestines simple, folded
into three segments. Specimens dioecious;
no evidence of hermaphroditism or oral
brooding.
Habitat: Engybenthic; continental slope
between approximately 200 and 1200
meters.
Key to Species of Epigonvs
la Opercular spine weak, poorly ossified, or
absent (opercular spine refers to the
ventralniost reinforced projection from the
posterodorsal edge of the opercle) 2
b Opercular spine pungent, bony 7
2a Lateral line scales 46-51; tongue eden-
tulous or bearing scattered tooth patches
3
b Lateral line scales 33-36; tongue cov-
ered with tooth patches (Fig. lA)
E. oligolepis
3a Gill rakers 23-34; premaxillary teeth short,
conical or villifonn, not visible when
mouth closed 4
b Gill rakers 17-21; premaxillary teetli
elongate, thin, inwardly recurved, visible
when mouth closed E. macrops
4a Pyloric caeca 7-14; first dorsal fin VII,
rarely VIII; vertebral count 10 + 15;
specimens not exceeding 220 mm SL 5
b Pyloric caeca 21-34; first dorsal fin VIII,
rarely VII (DiVIII often small or rudi-
mentary); vertebral count 11 -|- 14;
specimens to 550 mm SL E. telescopus
5a Body long, shallow; depth 15.8-23.6% SL;
peduncle length 25.4-32.2% SL; caudal
peduncle ring absent 6
b Body short, deep; depth 22.4-29.6% SL;
peduncle length 22.0-26.3% SL; caudal
peduncle ring present on specimens
shorter than 110-120 mm SL (Fig. IB)
E. pandionis
6a Gill rakers 28-34; pyloric caeca 10-14;
head length 31.2-38.6% SL; 2 pterygio-
phores between neural spines 9 and 10,
rarely 1 E. denticulatus
b Gill rakers 25-26, pyloric caeca 7-8;
head length 30.0-34.0% SL; 1 pterygio-
phore between neural spines 9 and 10
E. fiagilis
7a Body moderate to deep, 20.0-32.0% SL;
dorsal fins VII-1,9, rarely VII-1,10; gill
rakers 26-35 8
b Body shallow, 14.0-19.5% SL; dorsal fins
VII-1,10, rarely VII-1,9; gill rakers 22-27
— . E. occidentalis
8a Gill rakers of lower arch simple, awl-
shaped 9
b Gill rakers of lower arch pectinate ( Fig.
IC) - E. pectinifer
9a Tongue edentulous 10
h Tongue covered with tooth patches
E. trewavasac
10a Head length 28.0-36.6% SL; head height
14.7-18.8% SL; gill filaments moderate
or short 11
b Head length 36.8-41.9% SL; head height
18.9-21.1% SL; gill filaments long
,_ E. crassicaiidiis
11a Fin spines long, DJ 14.8-18.7% SL,
All 13.0-20.8% SL; interorbital width
8.7-10.2% SL; eyes large, 40.0-51.1% HL
E. leinmen
b Fin spines moderate, D2I 10.0-12.6% SL,
All 9.2-13.3% SL; interorbital width
6.5-8.2% SL; eyes moderate to small,
37.4-42.2% HL E. robustus
Epigonus telescopus (Risso, 1810)
Figure 2
Pomatomus telescopus Risso, 1810: 301, plate IX,
fig. 31 (original description; Nice; holotype
examined, MNHN B862); Lowe, 1841: 173;
Capello, 1868: 160; Moreau, 1881: 386, fig.
125; Vaillant (in part), 1888: 376.
Epigonus macwphthalmus Rafinesque, 1810: 64
( original description; no type locality; holotype
lost).
Pomatomus telescopium Cuvier, 1828: 171 (in-
correct emendation of Pomatomus telescopus
Risso, 1810); Valenciennes, 1830: 495;
Valenciennes, 1837-1844: 6, plate I; Giinther,
1859: 250; Cocco, 1885: 85; Holt and Calder-
wood, 1895: 405, plate LXIl.
Pomatomus cuvieri Cocco, 1829: 143 (original
description; seas of Messina; holotype not
examined ) .
Pomatotnus cuvicrii \'alenciennes, 1830: 501 (in-
correct emendation of Pomatomus cuvieri
Cocco, 1829).
?Pomatomichthys constanciae Giglioli, 1880: 20
( original description; Straits of Messina; holo-
type not examined, MZF 3089); Goode and
Bean, 1896: 234.
Epigonus telescopus Goode and Bean, 1896: 232;
Cligny, 1903: 9; Barnard, 1927: 523; Gall,
1931: 1, fig. 1; Fowler, 1936: 736, fig. 326;
Smith, 1949b: 206, fig. 474.
Scepterias lenimen, Whitley ( in part ) ( not Whit-
ley, 1935), 1968: 56.
Diagnosis. E. telescopus is the largest
species of the genus, growing to over 550
mm SL. Specimens are characterized by
21-34 pyloric caeca and eight first dorsal
EriaoNvs Systematics • Mayer 153
B
j^i^'
^^.^fffi^^'^h'"**'^^'
Figure 1. A. Tongue of E. oligolepis. Stippled areas
indicate tooth patches; shape and size of tooth patches
may vary among specimens. B. Caudal peduncle of
young E. pandionis showing anterior ring and posterior
band. C. Gill raker of E. pectinifer showing nub-like
processes.
fin spines. The opercular spine is blunt and
poorly ossified and distinguishes the species
from E. occidentalis, E. trewavosae, E.
pectinifer, E. rohiistus, E. lenimen, and E.
crassicaudus, which have pungent oper-
Y12yc
cular spines. Unlike remaining congeners,
E. telescopus possesses 11 + 14 vertebrae.
Description. Meristic data presented in
Table 1; regression data for morpho metric
traits presented in Table 2.
Body thickset, shortened; anterodorsal
profile slightly convex, rising most steeply
from tip of snout to interorbital region;
body moderate to deep, 21.2-26.3%^ SL;
caudal peduncle short, 19.0-26.5% SL.
Head moderate to deep, height 13.3-
SL; length 30..5-37.9% SL; snout
blunt; angle of gape moderate to large;
lower jaw equalling or protruding slightly
beyond upper jaw. Maxilla rarely exceed-
ing %-% eye length, posterior margin of
maxilla broad, posteriormost point near
ventral surface of bone; maxillae of large
specimens scaled. Eye round, 49.5-58.9%
HL; circumorbital tissues scaled, scale
pockets particularly apparent in large spec-
imens; anterodorsal rim of orbit projecting
into profile in small forms, reaching profile
in larger forms; interorbital width 9.0-
10.9% SL.
Dentition variable with age (see Onto-
genetic change); premaxillae, mandibles,
vomer, and palatines dentigerous; tongue
edentulous.
Opercle bearing short, poorly ossified
spine ventral to 1-8 membranous or poorly
ossified spinelets; spine and spinelets sepa-
rated by shallow gap; spinelets occasionally
obscured by underlying membranes. Pre-
opercle variable with age; angle rounded,
slightly produced in specimens shorter than
Figure 2. Epigonus telescopus, 220.0 mm SL, ISH 70/63.
154 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
Table 1. Epigonus telescopus meristic data. X = mean; SD = stan-
dard DEVIATION; n = NUMBER OF SPECIMENS.
Range
SD
Pectoral fin rays 20.85 19-23 0.71 54
Gill rakers 24.40 23-26 0.85 52
Lateral line scales 48.14 46-50 1.09 50
Pyloric caeca 25.25 21-34 3.59 16
200 mm SL, broadly produced in larger
forms; minute serrations on angle and ven-
tral surface of bone, rarely along posterior
surface dorsal to angle; striations radiating
from inner edge of angle. Interopercles
and subopercles without stiiations, occa-
sionally bearing minute serrations on pos-
tero ventral surfaces. Gill rakers simple,
awl-like.
First dorsal fin VII (7), VIII (46),
eighth spine small or rudimentary, lack-
ing membranous connection to preceding
spines; second dorsal fin 1,9 (1), 1,10 (52),
1,11 (1); anal fin 11,9 (56); D,I long,
3.5-6.3% SL; DJ, All short, 5.3-9.5%,
5.7-10.6%o SL respectively; Pol moderate,
6.5-11.9% SL.
Vertebrae 11 + 14 (18); epipleural ribs
Table 2. Epigonus telescopus regression data, b = regression coeffi-
cient ± 95%o CONFIDENCE INTERVAL; a = Y INTERCEPT; n = NUMBER OF
specimens. All regressions on SL.
b
a
n
HL
0.
35
+
0.
01
1.
60
50
Body depth
0.
25
+
0.
01
-2,
43
48
Head height
0.
19
+
0.
00
0.
45
45
Eye diameter
0.
13
+
0.
01
6.
57
44
Snout length
0.
10
+
0.
00
-2.
19
49
Interorbital width
0.
10
+
0.
00
-0,
32
52
Maxillary length
0.
16
+
0.
00
0.
00
48
Lower jaw length
0.
19
+
0.
00
-0.
61
50
Caudal peduncle de
pth
NONLINEAR
Caudal peduncle length
0.
2 1
+
0.
01
3.
35
51
D2I
0.
06
+
0.
02
3.
89
1 1
All
0.
06
+
0.
01
4.
18
31
P2I
NONLINEAR
Ei'iaoNus Systematics • Mayer 155
Table 3. Ontogenetic changes in the dentition of E. telescopus.
A. PREMAXILLARY DENTITION
< 200 mm SL
Extent 1/2-2/3 of ventral
surfoce
Pottern
I row
200-400 mm SL
2/3-7/8 of ventrol
surface
1-2 irregular rows
tapering to I row
> 400 mm SL
2/3-7/8 of ventral
surface
Multiple irregular rows
B. MANDIBULAR DENTITION
Extent
< 150 mm SL
Entire coronoid surface
150-250 mm SL
Entire coronoid surface
> 250 mm SL
Entire coronoid surface
Pottern
row
2-3 irregular rows
tapering to |-2 rows
3, 4, or 5 irregular rows
C. VOMERINE DENTITION
Extent
< I 75 mm SL
Center of vomer
> I 75 mm SL
Entire face of vomer
Pattern
Scattered teeth in few
irregular rows
Numerous teeth in multiple
irregular rows
D. PALATINE DENTITION
Extent
< 150 mm SL
Length of ventral surface
> I 50 mm SL
Length of ventral surface
Pottern
1-2 irregular rows
tapering to I row
2-5 irregular rows
tapering to I row
7 (11), 8 (2), inserting on vertebrae 1-7
or 1-8 respectively; pleural ribs 9 (17),
inserting on vertebrae 3-11.
Large specimens black or brown-violet,
iridescent in life ( Risso, 1810; Steindachner,
1891; Dons, 1938). Color in alcohol vari-
able with mode of collection and preser-
vation; skin often abraded, revealing under-
lying white-orange tissue; scale pockets
mottled with black or brown, melanophores
more densely packed near caudal edges;
pigment darker in larger fish; skin oily,
cutaneous fat deposits adding rust-colored
tint; opercular area black. Guanine de-
posits occasionally occurring on opercular,
tlioracic, and abdominal regions; iris black
with silver highlights; branchial membranes
black; mouth darkening with age (see
Ontogenetic change ) .
Description based on 54 specimens 68.1-
553 mm SL.
Ontogenetic change. Several marked
ontogenetic changes occur in E. telescopus,
the most noticeable involving dentition
patterns. Tooth-bearing bones of young
specimens exhibit relatively prominent con-
ical teeth. Teeth become more numerous
witli growth but appear smaller and form
weak conical or villiform bands. As Table
3 illustrates, older specimens have more
complex tooth patches with larger numbers
of tooth rows.
156 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
Epigonus Systematics • Mayer 157
A second change involves oral pigmenta-
tion. Young individuals have white or pale
yellow mouths; melanin is present only in
the vicinity of the pharynx. By the time
specimens reach 175-225 mm SL, black
pigment extends anteriorly to cover the
entire tongue. Shortly thereafter, the palate
becomes totally blackened, and by 300
mm SL, the entire mouth is dark.
The above changes are associated with
alterations in intestinal length. Measure-
ments of fourteen specimens ranging from
90.7-553 mm SL indicate that intestinal
length increases from 66-737^ SL in small
specimens (90.7-128.5 mm) to 98-108%
SL in moderate-sized individuals (220-250
mm). Thereafter, intestines grow more
slowly, reaching 110-115% SL in the largest
specimens. The coincidence of rapid in-
testinal growth, dentition changes, and de-
velopment of oral pigment suggests that
E. telescopus modifies its feeding habits
with growth.
Distribution. E. telescopus has an anti-
tropical distribution in the Atlantic, oc-
curring from Iceland to the Canary Lslands
and reappearing along the western coast
of South Africa (Fig. 3). Specimens have
also been taken in the Subtropical Con-
vergence region east of New Zealand. The
species is well known in the western Med-
iterranean and has been captured once
off the eastern coast of North America. A
single specimen is known from shallow
water off Norway (Dons, 1938).
Adults are taken by bottom trawl or long-
line and are most abundant from 300 to
800 meters; however, specimens have been
captured from water as shallow as 75 to
80 meters to as deep as 1000 to 1200 meters.
Koefoed (1952) reports four pelagic ju-
veniles from the Azores; Bertolini (1933)
mentions the presence of juveniles in the
Tyrrhenian Sea.
Earlier workers reported the range of E.
telescopus to include St. Helena (Val-
enciennes, 1837-1844; Giinther, 1868;
Bauchot and Blanc, 1961), tropical west
Africa (Osorio, 1898; Poll, 1954; Bauchot
and Blanc, 1961), and the Indian Ocean
(Steindachner, 1907; Fowler, 1935). These
accounts are based on misidentified or
tenuously identified material. The .speci-
mens described by Giinther, Poll, and
Bauchot and Blanc are E. pandionis, while
that examined by Fowler is Scomhrops-
like. Valenciennes' identification is ba.sed
on an impublished description and figure
by a St. Helena resident and must be re-
garded with suspicion. Reports by Stein-
dachner and Osorio could not be evaluated,
because neither includes a description or
figure of the material studied.
Geographic variation. The scarcity of
material from South Africa and New
Zealand makes it difficult to judge the
degree to which Northern and Southern
Hemisphere populations of E. telescopus
have diverged. Comparisons of dorsal and
pectoral fins, pleural and epipleural ribs,
lateral line scale counts, gill rakers, and
pyloric caeca reveal no subspecific dif-
ferences (coefficients of difference ^ 0.44).
Moiphometric characters, on the other
hand, exhibit greater variability. Of eight
traits successfully analyzed, three are sig-
nificantly different at both the 95%, 98%,
and 99% levels of confidence (Table 4).
These differences suggest that northern
and southern populations represent gemi-
nate subspecies; however, additional ma-
terial must be collected, especially from the
Southern Hemisphere, before definitive
statements can be made on intraspecific
variability.
Ta.xonomic notes. Pomatomichthtjs con-
stanciae Giglioli, 1880 is pro\'isionaily con-
sidered a junior synonym of E. telescopus
on the basis of work by Tortonese and
Queirolo (1970). These authors re-exam-
ined and, for the first time, figured the
holotype of P. constanciae. The latter
species is known only from the type speci-
men. The original description (Giglioli,
1880) is incomplete; no adequate rede-
scription has ever been published.
Data from the papers mentioned above
indicate a similarit\' between P. constanciae
158 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
Table 4. Comparison of regression coefficients from Northern and Southern
Hemisphere populations of E. telescopus. Data evaluated at the 95%, 98%,
and 99% levels of confidence. df = degrees of freedom; nb = regression
coefficients of Northern Hemisphere specimens; Sb = regression coefficients
OF Southern Hemisphere specimens; SD = significant difference between tab-
ular AND calculated VALUES OF t; t ^ CALCtJLATED VALUES OF t.
Significance
irt
CO
05
Nb
Sb
DF
t
a>
CT>
05
HL
0.
35
0
36
46
1. 54
Body depth
0.
25
0
24
44
0. 82
Head height
0.
20
0
18
41
3. 88
SD
SD
SD
Eye diameter
0.
13
0
12
40
1. 75
Snout length
0.
09
0
10
45
3. 19
SD
SD
SD
Interorbital width
0.
10
0
10
48
1. 05
Maxillary length
0
15
0
. 16
12
0 . 16
Lower jaw len
gth
0
19
0
. 19
14
0. 71
Caudal pedunc
depth
le
NONLINEAR
Caudal pedunc
length
le
0
21
0
. 22
47
0. 58
^2^
INSUFFICIENT
DATA
All
0
06
0
. 04
23
3. 80
SD
SD
SD
^2^
NONLINEAR
Table 5. Comparison of dorsal and pectoral fin counts from E. telescopus, P.
CONSTANCIAE, AND E. TREWAVASAE. DaTA FOR P. CONSTANCIAE FROM GiGLIOLI (1880)
AND TORTONESE AND QuEIROLO (1970); REMAINING DATA FROM PRESENT STUDY.
E. telescopus P. constanciae E. trewavasae
VII VII
I, 9.
1,9 rarely
I, 10
18 16-18
First
VIII,
dorsal
rarely
fin
VII
Second
I, 10,
dors a 1
r ar ely
fin
I, 9
Pectoral
fin
19 - 2:
Ei'iGONus Systematics • Mayer 159
Figure 4. Epigonus macrops, 154.6 mm SL, USNM 207679.
and E. telescopus but also suggest an af-
finity between P. constanciae and E. tre-
wavasae Poll, 1954. As is shown in Table
5, dorsal and pectoral fin counts fall within
the range of E. treicavasae rather than E.
telescopus. Tortonese and Queirolo's figure
similarly shows the holotype to possess a
sharp opercular spine, short DJ, and long
PJ — all characteristics of E. treicavasae.
Mensural data fail to differentiate P.
constanciae from either species. Unlike E.
treicavasae but like E. telescopus, the holo-
type lacks lingual teeth (Giglioli, 1880).
In view of the uncertainty surrounding
P. constanciae, a closer study of this form
must be undertaken. The problem is all
the more pressing, because E. treicavasae
is recorded from the Mediterranean for the
first time in this paper.
Common names. Comprehensive lists of
common names for E. telescopus are pro-
vided by Doderlein (1889), Nobre (1935),
and Bini (1968). Three names not re-
corded in these works are "Mejluza" — Gran
Canaria ( Steindachner, 1891), "Devil-fish"
— North Sea area (Ehrenbaum, 1928), and
"Big-eyed cardinal fish" — New Zealand
( Anonymous, 1961 ) .
Epigonus macrops (Brauer, 1906)
Figure 4
Oxijdon macrops Brauer, 1906: 288, fig. 172
(original description; Indian Ocean, land-locked
sea on west coast of Sumatra, VALDIVIA Sta.
186, 03°21'01"S, 101°11'05"E, 903 m; syntype
examined, ZMB 17678); Weber and de Beau-
fort, 1929: 351, fig. 81; Nomian, 1939: 60.
Diagnosis. E. macrops may be distin-
guished from all congeners by its low gill
raker counts ( 17-21 ) . It is further char-
acterized by eight fully developed first
dorsal fin spines and eight pyloric caeca,
one of which may function as a lumin-
escent organ.
Description. Meristic values presented
in Table 6; regression data for morpho-
metric traits presented in Table 7.
Body elongate; anterodorsal profile rising
steeply to occipital area; thereafter, weakly
convex, almost horizontal to first dorsal fin;
body depth 19.7-24.1% SL; caudal pe-
duncle length 22.0-26.7% SL.
Head length 34.1-38.5% SL; head licight
17.2-21.9% SL; snout blunt; angle of gape
large; lower jaw protruding beyond upper
jaw. Maxilla rarely exceeding Vs-% eye
length; posterior margin of maxilla broad,
bearing posteriormost point at ventral sur-
face of bone. Eye round to oval, 39.7-
48.3%' HL; anterodorsal rim of orbit pro-
jecting strongly into dorsal profile; inter-
orbital region wide, 9.5-11.7% SL.
Teeth conical, frequently recurved. Pre-
maxillary and mandibular teeth prominent,
needle-like, arranged in single row along
length of jaws; mandibular teeth occa-
sionally forming double row near sym-
physis; vomerine teeth few, moderate,
arranged in 2-4 irregular rows or in a
triangular or diamond-shaped patch; pala-
tin(\s bearing 2-6 teeth, arranged in single
row covering anterior half or second quar-
ter of bone; tongue edentulous.
Opercular spine short, weak, bony, ven-
160 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
Table 6. Epigonus macrops mebistic data. X = mean; SD = standard
DEVIATION; n = NLTNIBER OF SPECIMENS.
X
Range
SD
Pectoral fin rays 18.87 18-19 0.35 30
Gill rakers 18.63 17-21 0.87 32
Lateral line scales 48.61 46-50 0.83 28
Pyloric caeca 8.00 8 0.00 15
tral to 3-10 spinelets; spine and spinelets
separated by shallow, occasionally narrow
gap. Preopercular angle weakly produced,
rounded, serrate; serrations occasionally ex-
tending to posterior and ventral surfaces of
bone, rarely absent; striations radiating
from inner edge of angle. Subopercle and
interopercle generally serrated, occasionally
striated. Gill rakers short, awl-like.
First dorsal fin VII (1), VIII (29);
second dorsal fin 1,9 (1), 1,10 (31); anal
fin 11,9 (30), 11,10 (1). D^I, DJ, All
short, equalhng 1.2-2.9%, 5.3-7.7%, 5.9-
9.9% SL respectively; PJ moderate, 11.7-
14.1% SL.
Vertebrae 10 + 15 (25); epipleural ribs
6 (23), inserting on vertebrae 1-6; pleural
ribs 8 (24), inserting on vertebrae 3-10.
Table 7. Epigonus macrops regression data, b = regression coeffi-
cient ± 95% confidence interval; a = Y intercept; n =: number of
specimens. All regressions on SL.
b
a
n
HL
0.
35
+
0.
02
1
± •
48
26
Body depth
0,
22
+
0.
02
0.
51
31
Head height
0.
18
+
0.
02
1.
74
19
Eye diameter
0,
14
+
0.
02
3.
41
29
Snout length
0.
08
+
0.
01
-0.
18
22
Interorbital wi
dth
0,
11
+
0.
01
0.
36
30
Maxillary leng
th
0.
14
+
0.
01
1.
31
22
Lower jaw leu
gth
0,
18
+
0.
01
1.
18
31
Caudal peduncl
e dep
th
0.
12
+
0.
01
- 1.
53
29
Caudal peduncl
e len
gth
0.
24
+
0.
02
0.
70
30
D2I
0.
02
+
0.
02
7.
05
13
All
0.
04
+
0.
01
4.
73
22
P2I
0.
13
+
0.
02
0.
49
16
Ei'icoNus Systematics • Mayer
161
Figure 5. Caudal peduncle of young E. macrops bear-
ing anterodorsally canted ring.
Specimens probably black in life. Color
in alcohol variable with preservation; scale
pockets covered with black melanophores
near posterior edges; skin trecjiiently
abraded, revealing pink-yellow muscnla-
ture; opercular bones transparent, colored
black by underlying branchial membranes;
iris black; mouth black in adults. Young
bearing anterodorsally canted caudal pe-
duncle ring (see Ontogenetic change).
First pyloric caecum modified into lumin-
escent organ ( see Remarks ) .
Description based on 32 specimens 77.8-
206.0 mm SL.
Ontogenetic change. The transition from
juvenile to adult in E. macrops is marked
by changes in pigmentation and body
shape. Pelagic juveniles 15-37.9 mm SL
and young demersal forms 77.8-79.8 mm
SL bear a thin, black, anterodorsally tilted
ring circling the center portion of the
caudal peduncle (Fig. 5). Specimens
larger than 90 mm SL lack this marking.
Melanophores forming the rings are deeply
embedded in the peduncle musculature
and cannot be obliterated by abrading the
surface of the fish.
Adult E. macrops arc characterized by
black oral and branchial membranes. Al-
though these areas are colorless or poorly
pigmented in specimens smaller than 40
mm SL, the former surfaces darken and the
latter become covered with brown melano-
phores by the time fish reach 80 nun SL.
Juvenile E. macrops appear longer
and shallower than adults. Ratio-on-size
diagrams for interorbital width (i.e., in-
terorbital width/ SL vs. SL) indicate al-
lometric growth takes place in small
specimens. Similar statements are probably-
true for head height, eye length, and
caudal peduncle measurements but could
not be tested because of damage to juvenile
specimens.
DistriJnition. E. macrops adults are taken
exclusively by bottom trawls between 550
and 1100 meters in the Lidian Ocean,
Gulf of Mexico, Caribbean Sea, and West-
ern Atlantic. Specimens are most abundant
between 640 and 920 meters. Pelagic ju-
veniles are known from the Caribbean at
depths of 120 to 550 meters (Fig. 6).
GcograpJiic variation. No investigation
made because of inadec^uate Indian Ocean
samples.
Taxonomic notes. Brauer's description of
Oxyodon macrops ( 1906 ) is based on two
syntypes from the eastern Indian Ocean
( 172 and 212 mm total length ) . Of these,
only the larger is in the Zoologisches Mu-
seum der Humboldt Universitiit; the smal-
ler has been lost. The misplaced type may
have been deposited in the Zoologisches
Institut der Universitiit Leipzig and may
reappear when portions of this collection,
presently stored in Berlin, are sorted and
catalogued (Karrer, personal communica-
tion ) .
Remarks. Specimens of E. macrops bear
eight pyloric caeca; one of these appears
modified into a bioluminescent organ. The
luminescent caecum arises from the mid-
ventral surface of the pylorus just before
the duodenum and main body of pyloric
appendages (Fig. 7). It extends ventrally
until it reaches the floor of the abdominal
cavity, bends anteriorly and inserts into a
pouch formed by the black peritoneal
lining of the body cavity. At the posterior
edge of the pelvic girdle, the caecal pouch
lies over a thin, translucent portion of the
body wall which may function as a biolu-
minescent window. Externally the biolumi-
nescent window is covered by a single
large scale. The caecal pouch is lined with
silver or silver-gray pigment. Guanine
deposits appear most concentrated anter-
odorsally.
.\lthouah there is no direct evidence to
162 Bulletin Museum of Coiiiparative Zoology, Vol. 146, No. 3
m
in
^—
V
E
T3
O
<n
3
CO
SI
o
E
0}
>
3
■D
OS
TO
(0
<0
E
(U
■D
H—
o
"5
CO
JC
"cO
3
T3
>
T3
C
If)
<a
•
c
fl)
>
3
to
Q.
O
O
*^
TO
o
m
<n
E
CD
Q.
uj
O
.,
o
3
TO
c
r
o
3
TO
3
■n
■^-
■^
b
o
c
(D O
U. TO
Epiconus Systematics • Mayer 163
Figure 7. Luminescent organ of E. macrops. BW, body wall; D, duodenum; LPC, luminescent pyloric caecum;
LW, luminescent window; PC, nonluminescent pyloric caeca; PER, peritoneum; R, reflector; S, stomach.
support tlie claim that E. macrops is lu-
minescent, the modifications described
above are similar to those found in several
luminescent perciforms. Pernpheris klun-
zingeri and Parapriacanthus ransonneti
(Pempheridae) have luminescent organs
embedded in the thoracic ventral muscula-
ture formed from, or directly associated
with, the first pair of pyloric caeca ( Haneda
et al., 1966). Luminescent shallow-water
apog(jnids such as Apogon ellioti and
Siphamia nwiirnai also have luminescent
organs associated with the alimentary canal.
In both of the latter forms, anal and/ or
thoracic organs are connected by duct to
the intestine. As in E. nuicrops, tissue
above the luminescent structures may serve
as a reflector (Iwai, 1959; Haneda et al.,
1966).
Common names. None.
Epigonus pandionis (Goode
and Bean, 1881)
Figure 8
Apogon pandionis Goode and Bean, 1881: 160
( original description; off entrance to Cliesa-
peake Bay; holotype examined, USNM 26228);
Jordan and Gilbert, 1882: 564.
Figure 8. Epigonus pandionis, 141.7 mm SL, TABL uncatalogued.
164 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
Table 8. Epigonus pandionis meristic data. X = mean; SD = stan-
dard deviation; n = number of specimens.
X
Range
SD
Pectoral fin r ays
Gill rakers
Lateral line scales
Pyloric caeca
17. 81
17- 19
0. 57
97
27. 84
26- 30
0. 88
101
47. 63
46- 49
0. 66
81
10. 81
10- 13
0. 74
72
Glossamia pandionis Goode and Bean, 1896: 231.
Xystramia pandionis Jordan, 1917: 46.
Epigonus telescopus. Poll (not Risso, 1810),
1954: 89, fig. 26; Bauchot and Blanc (in part),
1961: 70.
Diagnosis. E. pandionis is the most ro-
bust species of the genus. Specimens
shorter than 110-125 mm SL are dis-
tinguished by a posterodorsally canted ring
circhng the caudal peduncle.
E. pandionis differs from E. macrops and
E. telescopus by bearing VII (rarely VIII)
spines in the first dorsal fin and 10-13
pyloric caeca. It is unlike E. oligolepis
because it has 46-49 lateral line scales and
may be distinguished from E. treioavasae,
E. pectinifer, E. robustus, E. lenimen, E.
crassicaudus, and E. occidentalis because
it lacks a pungent, bony opercular spine.
E. pandionis most closely resembles E.
fragilis and E. denticulatus but is differ-
entiated by its short caudal peduncle ( 22.0-
26.3% SL) and deep body (22.4-29.6%
SL). It further differs from E. dejiticulatiis
by exhibiting gill raker counts of 26—30
and a single basal pterygiophore between
neural spines 9 and 10.
Description. Meristic values presented in
Table 8; regression data for morphometric
traits presented in Table 9.
Body shortened, robust; anterodorsal pro-
file convex, particularly between occiput
and first dorsal fin; body deep, 22.4-29.6%
SL; caudal peduncle short, broad, length
22.0-26.3% SL.
Head length 33.0-39.0% SL; head height
19.0-22.2% SL; snout blunt; angle of gape
large; upper jaw subequal to lower jaw.
Maxilla reaching %-y-2 eye length; pos-
terior margin broad, posteriormost point at
ventral edge of bone. Eye round or slightly
oval, 37.4-48.7% HL; anterodorsal rim of
orbit projecting into profile in smaller
specimens, reaching profile in larger forms;
interorbital region wide, 9.2-11.5% SL.
Dentition variable with age (see Onto-
genetic change); premaxillae, mandibles,
vomer, and palatines dentigerous, bearing
conical, occasionally recurved teeth; tongue
edentulous.
Opercular spine short, horny, ventral to
2-5 (usually 3-4) poorly ossified spinelets;
spine separated from spinelets by gap;
spinelets occasionally obscured by un-
derlying membranes. Preopercular angle
broad, rounded, moderately produced;
striations radiating from inner ridge to
edges of angle; serrations along posterior
and ventral surfaces of bone. Subopercle
and interopercle bearing scattered serra-
tions. Gill rakers awl-like.
First dorsal fin VII (95), VIII (5); sec-
ond dorsal fin 1,9 (2), 1,10 (97), 1,11 (1),
11,10 (1); anal fin 11,8 (2), 11,9 (96), 11,10
(1); DJ long, 3.6-8.1% SL; DJ, All, PJ
short, equalling 5.0-8.6%, 5.0-8.7%, 8.9-
12.7% SL respectively.
Vertebrae 10 + 15 (36); epipleural ribs
6 ( 25 ) , 7 ( 1 ) , inserting on vertebrae 1-6
and 1-7 respectively; pleural ribs 8 (36),
inserting on vertebrae 3-10.
Epigonus Systematics • Mayer 165
Table 9. Epigonus pandionis regression data, b = regression coefft-
CIENT ± 95% CONFIDENCE INTERVAL; a = Y INTERCEIT; 11 = NUMBER OF
SPECIMENS. All regressions ON SL.
b
a
n
HL
0.
36
+
0.
01
0.
04
77
Body depth
0.
29
+
0.
01
-2.
64
75
Head height
0.
21
+
0.
01
-0.
30
67
Eye diameter
0.
16
+
0.
01
-0.
15
80
Snout length
0.
08
+
0.
00
0.
05
73
I n t e r 0 r b i t a 1 width
0.
1 1
+
0.
00
0.
33
74
Maxillary length
0.
17
+
0.
01
-0.
35
74
Lower jaw length
0.
19
+
0.
00
-0,
01
78
Caudal peduncle dep
th
0.
12
+
0.
00
-0.
61
80
Caudal peduncle len
gth
0.
24
+
0.
01
0.
54
81
D2I
0.
05
+
0.
01
1.
69
46
All
0.
05
+
0.
01
2.
11
56
P2 I
0.
10
+
0.
01
0.
71
75
Pigmentation variable with age (see
Ontogenetic change ) ; scale pockets mottled
with black; fin membranes black; opercular
region of adults black-slate gray; mouth
primarily light; iris black. Specimens fre-
quently abraded, underlying tissue pale
yellow-rust brown; guanine deposits rare,
if present occurring on opercular complex,
isthmus, thorax, or abdomen; silvered forms
generally from old collections.
Descriptions based on 104 specimens
45.7-194.0 mm SL.
Onto<ienetic chan2,e. Maturation in E.
pandionis is accompanied by changes in
pigmentation and dentition. The most
striking transformation involves caudal pe-
duncle markings. Specimens smaller than S5
mm SL bear a thin, black, posterodorsally
sloped ring circling the central portion of
the caudal peduncle. Melanophores form-
ing the ring are deeply embedded in pe-
duncle musculature and are not easily
abraded. A broader, more superficial band
of pigment circles the caudal peduncle at
the base of the caudal fin (Fig. IB). As
specimens grow beyond 85 mm, the rings
become fainter and begin to disappear.
Fish larger than 110 mm SL may com-
pletely lack peduncle markings, and by
125 mm SL, rings are absent from \irtually
all specimens. Since E. pandionis becomes
sexually mature at approximately 110 mm
SL, altered markings may reflect changes
in habit or behavior associated with repro-
ductive individuals.
Gill rakers and branchial membranes are
converted from pale yellow to black. Spec-
imens smaller than 55 mm SL bear scat-
tered black melanophores on gill rakers
but lack opercular pigmentation. B\- 60
mm SL rakers have become totalK' dark,
and traces of melanin ha\'e appeared on
membranes lining the opercle. Pigment
becomes denser with growth and spreads
ventrally. By 100 mm SL the opercle is
completely lined with dark tissue. Since
166 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
Ei'iGONus Systematics • Mayer 167
opercular bones are translucent, the process
appears outwardly as a darkening of the
opercle.
Ontogenetic changes in dentition involve
the production of increasingly complex
tooth patches. Specimens smaller than 80
mm SL bear single rows of teeth on the
premaxillae and palatines. Mandibular teeth
are arranged in patterns analogous to those
found on the premaxillae or in double rows
that taper to a single row posteriorly.
Vomerine teeth occur in 1-2 chevron-
shaped clumps. As growth takes place,
teeth are added to all dentigerous surfaces.
Large specimens (>130 mm SL) have as
many as 3-4 tooth rows on palatines and
anterior segments of dentaries and premax-
illae. Vomerine teeth may become suf-
ficiently numerous to cover the entire face
of the bone.
Distribution. E. pandionis is amphi-At-
lantic, occurring primarily in the Caribbean,
Gulf of Mexico, and Gulf of Guinea (Fig.
9). The species has been taken as far
north as New Jersey and as far south as
French Guiana in the western Atlantic.
It occurs between Portuguese Guinea and
Angola in the eastern Atlantic. Adults are
captured exclusively by bottom trawls be-
tween 210 and 600 meters. American forms
are most numerous from 300 to 500 meters,
while African populations are most abun-
dant between 260 and 450 meters. A single
pelagic juvenile (35.5 mm SL, MCZ 48839)
was taken at 275 to 300 meters in the
Caribbean.
Geo<:,raphic variation. Statistical analyses
provide conflicting assessments of the
similarity of African and American popula-
tions. Meristic characters reveal little vari-
ability. Coefficients of difference calculated
for standard counts are always less than
or equal to 0.49 — far below conventional
levels of subspecies recognition. Mensural
data, on the other hand, suggest there are
considerable differences between the pop-
ulations. Of thirteen traits analyzed, seven
separate eastern and western populations
at the 95% level of confidence, five separ-
ate them at the 98% level, and two separate
them at the 99% level (Table 10).
A closer examination of the characters
exhibiting signilicant differences reveals
that regression coefficients of American
E. pandionis are always greater than those
of African forms. Since regression coef-
ficients are a measure of relative growth,
observed intraspecific variation may reflect
environmental factors.
Water temperature is a major parameter
determining growth rates in fishes. If other
factors are conti'olled, rates of growth in-
crease proportionally with temperature
(Brown, 1957: 391). With this in mind, it
is interesting that temperatures are gener-
ally higher and superficial warm-water
layers thicker in the western tropical At-
lantic (Ekman, 1953). At 300 meters Gulf
of Mexico and Caribbean temperatures vary
from 10 to 18° C while west African tem-
peratures range between 9 and 11° C. At
500 meters the difference is slightly less
pronounced — 8-13° C as opposed to 6-8°
C (from temperatin-e profiles in Fuglister,
1960; Wiist, 1964; and Nowlin and
McLellan, 1967). One would therefore
expect western Atlantic E. pandionus to
grow more rapidly and exhibit larger re-
gression coefficients than eastern Atlantic
forms. In view of these findings, the two
morphs are not considered to represent
separate subspecies.
Remarks. See E. trewavasae: Remarks
for discussion of E. pandionis .sensu Lozano
(1934), Navarro et al. (1943), and Maurin
(1968).
Specimens of doubtful identity. Five spec-
imens were examined that resembled E.
pandionis but could not, with certainty,
be placed in the species. Four were taken
in the Atlantic, the fifth in the Gulf of
Oman (see Mayer, 1972: Appendix II for
complete data). These fishes were not
considered when preparing the description
of E. pandionis, nor were they used in
morphometric, meristic, or distribution
analyses.
Tlie Atlantic specimens include three
168 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
Table 10. Comparison of regression coefficients from eastern and %vestern
Atlantic populations of E. pandionis. Data evaluated at the 95%, 98%, and
99% LE\rELS OF confidence. DF =: DEGREES OF FREEDOM; Eb = REGRESSION COEFFI-
CIENTS OF EASTERN ATLANTIC SPECIMENS; SD = SIGNIFICANT DIFFERENCE BETWEEN
TABULAR AND CALCULATED VALUES OF t; t=3 CALCULATED VALUES OF t; Wb = REGRES-
SION COEFFICIENTS OF WTESTERN ATLANTIC SPECIMENS.
Significance
Wb Eb DF t S S S
HL
0.
36
0.
37
60
0.
97
Body depth
0.
29
0.
29
71
0,
31
Head height
0.
22
0.
20
63
2.
41
SD
SD
Eye diameter
0.
17
0.
15
76
2.
14
SD
Snout length
0.
09
0.
08
69
3.
00
SD
SD
SD
Interorbital width
0.
11
0,
11
70
2.
13
SD
Maxillary leng
^th
0.
18
0.
16
70
3.
35
SD
SD
SD
Lower jaw len
gth
0.
20
0.
18
74
2.
63
SD
SD
SD
Caudal pedunc
depth
le
0.
13
0.
12
76
2.
39
SD
SD
Caudal pedunc
length
le
0.
23
0.
24
77
1.
16
D2I
0.
05
0.
06
32
1
83
All
0.
05
0.
05
52
0
91
P2I
0.
10
0.
10
71
0
83
fishes from St. Helena. The most recently for only the Caribbean form, which was
collected (UZM P45148) was incorrectly taken at relatively shallow depths. Exact
identified as E. telescopus by Banchot and determination of the variants' status must
Blanc (1961). The two older fonns (BMNH await the capture of additional material.
1868.3.11.14/15) are probably the fish The Indian Ocean form (BMNH
discussed by Giinther (1868). The re- 1889.4.15.24) is distinguished from E.
maining specimen (USNM 207703) was pandionis by its shallow body (22.5% SL),
taken in the Caribbean. narrow interorbital region (8.3% SL), den-
The four Atlantic individuals are basi- tigerous glossohyal, numerous weak oper-
cally similar to E. pandionis but exhibit cular spinelets, and elongate gill filaments,
shallower heads ( 17.4-19.8% SL), narrower The last trait suggests the fish may have
interorbital regions (8.6-9.4% SL), fewer inhabited an oxygen minimum layer. As
pyloric caeca (8-9), and fewer gill rakers with the Atlantic variants, additional ma-
( 25-27 ) . In these respects they resemble terial must be collected before the status of
E. fragilis. Little is known about the habits the form can be determined,
of the variants; station data are available Common names. None.
Epigonus Systematics • Mayer 169
Z^A
^
y
y
x»
Figure 10. Epigonus fragilis, HOLOTYPE, 89.1 mm SL, CM 3900/FMNH 55204 (from Jordan and Jordan, 1922).
Epigonus fragilis (Jordan and
Jordan, 1922)
Figure 10
Scepterias fragilis Jordan and Jordan, 1922: 45,
plate II, fig. 2 (original description; Honolulu
market; holotype examined, CM 3900/FMNH
55204).
?Hynnodus fragilis Pietschmann, 1930: 13.
Diagnosis. E. fragilis most closely re-
sembles E. pandionis but may be dis-
tinguished by its shallow body (18.8-21.1%
SL) and short, shallow head (length 31.7-
34.0% SL, height 16.0-17.4% SL). Unlike
£. pandionis, E. fragilis lacks peduncle
rings on specimens smaller than 100-120
mm SL.
In the past E. fragilis has been confused
with Hijnnodus atherinoides, a junior syn-
onym of E. occidentalis. E. fragilis may be
distinguished on the basis of body depth
( see above ) , pectoral fin counts ( 16-17 ) ,
and the absence of a pungent, bony oper-
cular spine. Weak opercular armor, to-
gether with second dorsal fin counts of 1,10
differentiate E. fragilis from E. treivavasae,
E. pectinifer, E. rohustus, E. lenimen, and
E. crassicaudus. Gill raker counts of 25-26
separate E. fragilis from all remaining con-
geners except E. telescopiis. E. fragilis may
be distinguished from the latter by the
presence of 7-8 pyloric caeca.
Description. E. fragilis is known from
only five specimens. Of these, the holotype
is of little descriptive value. The specimen
is severely dehydrated and has become
discolored, brittle, and shrunken. The fol-
lowing account is based primarily on two
recently captured specimens of E. fragilis
(LACM 32668-6 and USNM 207704) and
two forms collected by D. S. Jordan in 1921
(SU 23246). The latter are mentioned in
the original description of E. fragilis but
are not designated as types.
All meristic and mensural data are pre-
sented in the text. Detailed statistical
analyses were not undertaken because of
small sample size.
Body elongate; anterodorsal profile con-
vex, rising without interruption from tip
of snout to first dorsal fin. Body depth
18.8-21.1% SL; caudal peduncle length
25.4-26.9% SL.
Head short, 31.7-34.0% SL; head height
16.0-17.4% SL; snout blunt, 7.2-7.9% SL;
angle of gape moderate; jaws equal. Max-
illa reaching % eye length; posteriormost
point of maxilla at ventral edge of bone.
Eye round, 38.1—41.5% HL; anterodorsal
rim of orbit reaching profile; interorbital
width 8.8-9.4% SL.
Dentition variable with age. Teeth con-
ical; premaxillary teeth in irregular double
rows anteriorly, tapering to single row
posteriorly, occupying anterior %-% of
bone. Mandibular dentition more promin-
ent than that of premaxilla; teeth recurved,
occupying from % to entire length of
dentary, arranged in single or double rows
near symphysis and single row posteriorly.
Vomerine teeth recurved, arranged in oval
or diamond-shaped patch, covering entire
face of bone in adults. Palatine teeth
170 Bulletiti Museum of Comparative Zoology, Vol. 146, No. 3
Figure 11. Epigonus occidentalis, 152.7 mm SL, MCZ 48840.
medially recurved, arranged in single-triple
rows anteriorly, tapering to single row
posteriorly; tongue edentulous.
Opercular spine weak, ventral to 7-9
small serrae; angle of preopercle produced,
rounded, ornamented with striations and
weak serrations; subopercle and inter-
opercle unornamented. Gill rakers 25 (3),
26 (1), simple, awl-like. Pyloric caeca
7(1), 8(2).
First dorsal fin VII (4), VIII (1);
second dorsal fin 1,10 (5); anal fin 11,9
(5); pectoral fin 16 (1), 17 (3); DJ
moderate to long, 5.9-8.9% SL; DJ short,
6.9% SL; PJ long, 10.1-10.2% SL; All
broken.
Vertebrae 10 + 15 (4); epipleural ribs
not visible on radiographs; pleural ribs
8 (4), inserting on vertebrae 3-10. Pored
lateral line scales 49 ( 2 ) .
Color in alcohol yellow-brown; fin mem-
branes dark; iris silver-black; mouth light;
branchial membranes light, darkening with
age.
Distribution. E. fragilis is endemic to
the Hawaiian Islands (Fig. 12). The spe-
cies is demersal and has been taken between
120 and 125 meters.
Taxonomic notes. Six years after E.
fragilis was described, Fowler ( 1928 ) syn-
onymized the species with a second Ha-
waiian apogonid, Hijnnodus atherinoides
Gilbert, 1905. The synonymy achieved
moderate acceptance and appeared in sev-
eral publications (e.g., Matsubara, 1936;
Tinker, 1944; Gosline and Brock, 1960).
Fowler's conclusions were based on a
33-mm specimen (BPBM 3914) obtained
by the Tanager Expedition. The specimen
is in extremely poor condition. All colora-
tion has been lost, most of the muscle
tissue has decomposed, and much of the
skeleton has become decalcified. Although
it is impossible to identify the fish because
of its condition, the following traits suggest
it is neither E. fragilis nor H. atherinoides:
dorsal fin elements — VIII-1,8; anal fin ele-
ments— 11,6; vertebrae — 11 + 14. These
data differ from Fowler's report of VI-I,8
dorsal elements, no anal spines, and 7 anal
rays.
As was discussed in the diagnosis, E.
fragilis is distinct from H. atherinoides.
Fowler's synonymy appears to have been
based on inaccurate data taken from an
incorrectly identified fish.
Common names. None.
Epigonus occidentalis Goode
and Bean, 1896
Figure 11
Epigonus occidentalis Goode and Bean, 1896: 233,
plate LXVI, fig. 236 (original description;
Steamer BLAKE, off Barbados, 237 fms.; holo-
type examined, MCZ 28032 ) .
Hijnnodus atherinoides Gilbert, 1905: 618, plate
79 (original description; ALBATROSS Sta.
3867, Pailolo Channel, Hawaii, 284-290 fms.;
holotype examined, USNM 51601); Jordan
and Jordan, 1922: 44; Fowler and Bean, 1930:
121.
Hijnnodus megalops Smith and Radcliffe, 1912
{in Radcliffe, 1912): 445, plate 38, fig. 3
(original description; ALBATROSS Sta. 5388,
12°51'30"N, 123°26T5"E, between Bnrias and
Luzon, Philippines, 226 fms.; holotype ex-
amined, USNM 70255).
Ei'iGONus Systematics • Mayer 171
Table 11. Epigonus occidentalis meristic data. X = mean; SD
STANDARP nEVIMION; n = NUMBER OF SPECIMENS.
X
Range
SD
Pectoral fin rays
Gill r aker s
Lateral line scales
Pyloric caeca
20.
21
19-2 1
0. 59
56
24.
68
22-27
1. 08
60
48.
15
46-51
0. 97
46
9.
27
8-13
1. 05
45
Table 12. Epigonus occidentalis regression data, b = regression
COEFFICIENT ± 95% CONFIDENCE INTERVAL; a = Y INTERCEPT; n = NUMBER
OF SPECIMENS. AlL REGRESSIONS ON SL.
b
a
n
HL
0.
34
+
0.
02
0. 72
48
Body depth
0,
19
+
0.
02
-1. 72
48
Head height
0.
15
+
0.
01
0. 53
49
Eye diameter
0.
16
+
0.
01
0, 66
49
Snout length
0.
08
+
0.
00
0. 06
49
Interorbital width
0.
08
+
0,
01
0. 83
39
Maxillary length
0.
13
+
0,
01
0. 88
51
Lower jaw length
0,
15
+
0.
01
1. 26
51
Caudal peduncle de
pth
0,
10
+
0.
01
-0. 90
54
Caudal peduncle len
igth
0.
23
+
0.
01
1. 50
53
D2 I
0.
05
+
0.
00
1. 42
34
All
0.
05
+
0.
01
2. 18
42
P2I
0.
09
+
0.
01
0. 67
47
Diagnosis. E. occidentalis is distin-
guished from all other congeners by the
combination of shallow body depth (14.1-
19.57t SL), reduced gill raker counts (22-
27), and the presence of a pungent, bony
opercular .spine. It is frequently confused
with E. denticulatus.
Description. Meristic values presented
in Table 11; regression data for morpho-
metric traits presented in Table 12.
Body elongate, cigar-shaped; anterodor-
sal profile weakly convex, flattened, rising
gradually from tip of snout to interorbital
region, leveling off toward occipital region,
and rising gradually to base of first dorsal
fin. Body depth 14.1-19.5% SL, body
172 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
width subequal to or greater than body
depth; caudal peduncle narrow, length
22.4-28.1% SL.
Head length 30.5-37.9% SL; head height
13.3-17.2% SL; angle of gape moderate to
small; lower jaw equalling or protruding
slightly beyond upper jaw. Maxilla reach-
ing Vi-% eye length; posterior margin
of maxilla moderate to narrow, posterior-
most point at ventral edge of bone. Eye
long, oval, 40.6-52.3% HL; anterodorsal
rim of orbit reaching or projecting into
dorsal profile; interorbital region narrow,
5.6-8.5% SL.
Teeth conical; premaxillary and man-
dibular teeth frequently recurved, arranged
in simple single row or single row widening
to double or triple rows near symphysis;
teeth covering % to entire length of pre-
maxilla and % to entire length of dentary;
vomerine teeth arranged in 1-4 irregular
rows; palatines rarely edentulous, teeth
1-10, arranged in single row, covering
anterior Vi-V^ of bone; tongue edentulous.
Opercular spine pungent, bony, ventral
to 1-3 poorly ossified spinelets; spine sep-
arated from spinelets by shallow indentation.
Preopercular angle produced, rounded or
pointed, bearing serrations and striations;
subopercle serrate, occasionally striate;
interopercle variable, frequently serrate.
Gill rakers short, awl-like.
First dorsal fin VII (59); second dorsal
fin 1,10 (59); anal fin 11,8 (1), 11,9 (59);
DJ, DJ, All, PJ short, equahing 1.1^.2%,
4.8-7.8%, 4.8-9.2%, and 8.0-11.3% SL re-
spectively.
Vertebrae 10 -I- 15 (35); epipleural ribs
6 (19), 7 (5), inserting on vertebrae 1-6
or 1-7 respectively; pleural ribs 7 (31),
8(1), inserting on vertebrae 2-9 or 3-9.
Color in alcohol variable with preser-
vation; skin frequently removed by trawl-
ing; underlying tissue pale yellow, yellow-
pink, occasionally marked with rust brown;
scale pockets and fin membranes black;
opercular area black-slate gray, occasion-
ally tinged with silver; lower jaw, bran-
chiostegal membranes, and thoracic and
abdominal regions occasionally silvered;
guanine most prevalent on specimens from
old collections. Mouth color variable with
age (see Ontogenetic change); iris and
branchial region dark.
Description based on 62 specimens 58.2-
178.9 mm SL.
Ontogenetic change. The most striking
age-related change in E. occidentalis is
the development of oral pigmentation. As
in E. telescopus and E. macro'ps, immature
forms bear pigmentless or slightly pig-
mented mouths, while adults have black-
ened oral membranes. Pigmentation first
appears in specimens 80-110 mm SL.
Melanophores develop just anterior to the
pharynx and spread rostrally, covering a
third of the roof and floor of the mouth and
half of the tongue by the time specimens
reach 115-130 mm SL. By 150 mm SL the
tongue is completely black, and by 175-
180 mm the entire mouth is dark. Branchial
membranes undergo an analogous trans-
formation before specimens reach 58 mm
SL.
A faint black ring circling the middle of
the caudal peduncle was observed on
three small E. occidentalis (< 65 mm SL).
Similar markings were absent from larger
individuals. The rings are reminiscent of
markings observed on young E. macrops
and E. pandionis and probably represent
a juvenile feature that is lost with growth.
Distribution. E. occidentalis has been
taken in the Caribbean, Gulf of Mexico,
and western tropical Atlantic. It is also
known from the Philippine and Hawaiian
Islands (Fig. 12). The species is caught
by bottom trawls between 360 and 735
meters. Adults are most abundant in the
Caribbean from 500 to 550 meters.
Geographic variation. E. occidentalis, as
here defined, includes two nominal species
— Hijnnodus atherinoides Gilbert, 1905 and
H. megalops Smith and Radcliffe, 1912.
The former originally represented a Ha-
waiian endemic; the latter represented a
Philippine form. In 1930 Fowler and Bean
synonymized the Pacific morphs. In the
Epigonus Systematics • Maxjer 173
Figure 12. Distribution of E. fragilis and E. occidentalis. Map A shows localities
in the Caribbean and Gulf of Mexico. Map B shows localities in the western
Pacific. J^ E. fragilis, individual haul of demersal adults; % E. occidentalis, indi-
vidual haul of demersal adults; cross-hatching indicates areas where E. occiden-
talis are frequently taken.
174 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
3 cm
Figure 13. Epigonus denticulatus, 115.1 mm SL, UMML 12463.
authors' opinions, characters separating the
two forms were "simply minor discrepancies
of portraiture and should never have been
credited as specific distinctions [p. 122] ."
Although descriptions and illustrations
of H. atherinoides and H. megalops suggest
a link with E. occidentalis, detailed com-
parisons of the three forms were never
made. To a large extent this was the re-
sult of inadequate sampling. Until the
initiation of the OREGON cruises in 1950,
few E. occidentalis were available for study.
Pacific forms are still poorly represented;
only seven specimens have been collected.
Reports of additional material by Fowler
(1928), Matsubara (1936), Smith ( 1949a,b,
1961), Kamohara (1952), and Moreland
( 1957) are based on misidentifications.
Comparisons of E. occidentalis and the
H. atherinoides-H . megalops complex pro-
vide no evidence to support their status
as separate species. Analyses of head
length, body depth, head height, eye di-
ameter, snout length, interorbital and max-
illary widths, caudal peduncle length and
depth, and All and PJ lengths reveal no
significant differences between the pop-
ulations at either the 95%, 98%, or 99%
levels of confidence. Meristic data also
show considerable overlap for most char-
acters; however, the coefficients of dif-
ference for pyloric caeca and gill raker
counts are above conventional levels of
subspecies recognition (1.68 and 1.99, re-
spectively). In addition, Atlantic and
Pacific populations may be distinguished
by minor qualitative characters such as:
(1) short, rounded preopercular angles
in Atlantic forms; longer, pointed
angles in Pacific specimens;
(2) fusion of uroneurals 1 and 2 in
Atlantic forms (based on 3 alizarin
preparations ) ; separate occurrence
in Pacific forms (based on 1 alizarin
preparation ) .
On the basis of the above information,
Atlantic and Pacific forms are placed in
the same species but considered members
of separate subspecies. Formal description
of the subspecies must await the capture
of additional Pacific specimens.
Remarks. A single unripe female E.
occidentalis (USNM 197353, 172.1 mm SL)
was found carrying small egg masses in the
anterior portion of its mouth (anterior to
the tongue and vomer). The masses con-
tained 125 oval eggs 0.40-0.55 mm in
diameter. The presence of eggs in the
mouth of an Epigonus is of interest, be-
cause several shallow-water apogonids ex-
hibit oral brooding. No such activity has
ever been reported for deep-sea forms.
Although it is difficult to say with
certainty, the E. occidentalis eggs are prob-
ably not incubating clutches, but rather
non-apogonid ova ingested during trawling.
Unlike the egg masses of typical oral
brooding apogonids, those found in E. oc-
cidentalis are broken, disrupted, and con-
tain very few eggs. An 84.9-mm specimen
of Cheilodipterus affinis was reported in-
Epigonus Systematics • Mayer 175
Table 13. Epigonus denticulatus meristic data. X = mean; SD
STANDARD DEVIATION; n = NUMBER OF SPECIMENS.
X Range SD n
Pectoral fin rays
19.
09
18-20
0. 56
54
Gill rakers
30.
98
28- 34
1. 10
58
Lateral line scales
48.
12
46- 49
0. 76
43
Pyloric caeca
11.
83
10- 14
0. 85
42
ciibating 21,000 eggs 0.35-0.4 mm in di-
ameter (Smith et al., 1971: 8-9). The ova
fully occupied the oral and branchial
chambers and extensively distended the
head. These conditions were not observed
in E. occidentalis.
It is possible that the eggs represent the
remnants of a larger mass that was spit
out and partially reingested. However,
were this the case, one might expect to
find eggs in the stomach (Sakomoto, 1930)
or gill rakers. No eggs were found in
either region.
Finally, Breder and Rosen (1966) state
that eggs of oral brooding apogonids are
lield together by fibers attaching to one
pole. The eggs of E. occidentalis are
loosely embedded in an open matrix of
fibers. Grape-like egg clusters character-
istic of Apogon semilineatus (Ebina, 1931:
20 ) were not observed.
Common names. None.
Epigonus denticulatus Dieuzeide, 1950
Figure 13
Pomatomus telescoptis, Vaillant (in part) (not
Risso, 1810), 1888: 376.
Scepterias lenimcn, Whitley (in part) (not Whit-
ley, 1935), 1935: 230; Whitley (in part),
1940: 420.
Epigonus atherinoides, Matsubara (not Gilbert,
1905), 1936: 120, fig. lA; Smith, 1961: 378,
fig. 3; Kamohara, 1952: 37, fig. 31.
Hynnodus atherinoides. Smith (not Gilbert, 1905),
1949a: 101; Smith, 1949b: 210, fig. 495A.
Epigonus denticulatus Dieuzeide, 1950: 89, figs.
1-2 (original description; Algerian Coast at
200-500 m; holotype not examined); Tortonese,
1952: 72, 1 fig.; Dieuzeide et al., 1953: 216,
2 figs.; Tortonese and Queirolo, 1970: 33,
fig. 6.
Diagnosis. E. denticulatus lacks a fully
ossified opercular spine, bearing instead
3-7 membranous projections. This feature
distinguishes it from E. occidentalis, E.
treivavasae, E. pectinifer, E. rohustus, E.
lenimen, and E. crassicaudtis, which have
pungent, bony opercular spines. E. denti-
culatus is differentiated from E. telescopus,
E. macrops, and E. fragilis by the presence
of 10-14 pyloric caeca and 28-34 gill
rakers. It differs from E. oligolepis by
bearing 46-51 lateral line scales. E. denti-
culatus closely resembles E. pandionis but
may be distinguished on the basis of the
former's shallow body (15.8-23.67r SL),
long caudal peduncle (25.9-32.2% SL),
and short DJ (2.4-3.7% SL).
Description. Meristic values presented
in Table 13; regression data for morpho-
metric traits presented in Table 14.
Body fusiform, slightly compressed; an-
terodorsal profile rising gradually above
snout, becoming steeper and slightly con-
vex over eyes, thereafter rising gradually
to first dorsal fin; body moderate to shal-
low, depth 15.8-23.6% SL; caudal peduncle
narrow, length 25.9-32.2%o SL.
Head moderate to short, 31.2-38.6% SL;
head height 16.0-19.8% SL; snout short,
blunt; angle of gap(> moderate to large;
lower jaw protruding slightly beyond up-
per jaw. Maxilla reaching %-% eye length,
176 Biilletifi Museum of Comparative Zoology, Vol. 146, No. 3
Table 14. Epigonus denticulatus regression data, b = regression
COEFFICIENT ± 95% CONFIDENCE INTERVAL; a = Y INTERCEPT; 11 = NUMBER
OF SPECIMENS. AlL REGRESSIONS ON SL.
HL
0.
32
+
0.
01
2.
88
57
Body depth
0.
25
+
0.
01
-4.
09
54
Head height
0.
16
+
0.
01
0.
86
56
Eye diameter
0.
14
+
0.
01
1.
39
58
Snout length
0.
07
+
0.
00
0.
33
56
Interorbital width
0.
09
+
0.
00
0.
37
55
Maxillary length
0.
14
+
0.
01
1.
37
56
Lower jaw length
0.
15
+
0.
01
1.
61
57
Caudal peduncle d
epth
0.
11
+
0.
, 01
-0.
78
57
Caudal peduncle li
ength
0.
28
+
0.
, 01
0.
61
57
D2I
0.
05
+
0.
, 01
2.
24
36
All
0.
, 06
+
0,
, 01
1.
, 43
40
P2I
0.
, 08
a.
0,
, 01
0.
, 92
41
posteriormost point near ventral surface
of bone. Eye round or slightly oval, 40.3-
48.0% HL; anterodorsal rim of orbit
reaching dorsal profile, projecting into pro-
file in smaller specimens; interorbital
width 8.2-10.4% SL.
Teeth small, conical, occasionally re-
curved; premaxilla bearing single row of
teeth along anterior Vs-% (usually %) of
bone. Mandibular teeth arranged along
length of dentary in irregular single row,
occasionally double near symphysis; larger
specimens with 3-4 rows near symphysis.
Vomerine teeth variable, arranged in 1-4
irregular rows. Palatine dentition occupy-
ing length of bone, arranged in simple
single row or double row tapering to single
row posteriorly; large specimens bearing
3-4 rows of teeth anteriorly. Tongue gen-
erally edentulous, rarely Ijearing isolated
tooth patches on glossohyal or edges of
tongue.
Opercle lacking bony spine, bearing in-
stead 3-7 (usually 5-6) jagged, mem-
branous projections; projections often ob-
scured by underlying tissues. Peropercular
angle produced, broadly rounded, striations
radiating from inner edge, angle occasion-
ally serrate; subopercle and interopercle
occasionally serrate. Gill rakers simple,
awl-like.
First dorsal fin VII (53); second dorsal
fin 1,9 (1), 1,10 (56), 10 (1); anal fin 11,8
(1), 11,9 (57). DJ moderate, 2.4-3.7%
SL; Dol, All, P,I short, 5.2-8.0%, 6.0-8.2%,
7.9-10.0% SL respectively.
Vertebrae 10 + 15 (44); epipleural
ribs 6 ( 32 ) , 7 ( 1 ) , inserting on vertebrae
1-6 or 1-7 respectively; pleural ribs 8 ( 44 ) ,
inserting on vertebrae 3-10.
Color in alcohol variable with preserva-
tion; skin frequently removed by trawling,
underlying tissue pink-brown or yellow;
scale pockets mottled with numerous
brown-l)lack melanophores, dorsal surfaces
of body and head more heavily pigmented.
Epigonus Systematics • Mayer 177
Guanine deposits frequently occurring on
gill cover, ventral surface of mandible,
isthmus, thoracic region, and abdomen to
anus; iris black; mouth light; branchial
region dark.
Description based on 58 specimens 57.0-
187.5 mm SL.
Ontogenetic change. Two young spec-
imens of E. dcnfictihifus (29.2 mm SL,
MCZ 48846, and 49.7 mm SL, MCZ 48847)
were examined in the course of this in-
vestigation. These specimens were taken
by midwater trawls made in the central
North Atlantic and Gulf of Mexico and
reveal that the life cycle of E. denticulatus
includes a pelagic juvenile stage.
The pelagic young resemble adults in
most respects. For example, the juveniles
bear diagnostic gill raker counts and
opercular ornamentation. However, slight
changes in body shape are associated with
growth. The 29.2 mm specimen has a more
shallow body, shorter head, narrower inter-
orbital region, and smaller eyes than
demersal adults. Similar trends are present
but less apparent in the larger juvenile.
Juvenile dentition patterns are basically
like those of adults but involve fewer and
relatively larger recurved teeth. Oral and
branchial regions are light in young speci-
mens. The latter areas darken with age.
Distribution. E. denticulatus is the only
cosmopolitan species in the genus (Fig.
14). Specimens have been taken from the
southwest coast of Japan, the Gulf of
Mexico, and the Caribbean. In addition,
the species occurs continuously from the
western Mediterranean, south along the
western coast of Africa to the tip of the
continent. It reappears south of the Great
Australian Bight and southeast of New
Zealand.
Adults are generally taken by bottom
trawls between 300 and 600 meters, al-
though specimens have been captured from
as shallow as 200 meters and as deep as 830
meters. Pelagic juveniles have been taken
by IKMT between 130 to 145 meters and
350 to 425 meters.
Geographic variation. E. denticuhitus
may be divided into North Atlantic, South-
ern Hemisphere, and Japanese populations.
North Atlantic forms include material from
the Mediterranean, northeast Atlantic,
Caribbean, and Gulf of Mexico. Southern
Hemisphere populations contain specimens
from the southeast Atlantic, Australia, and
New Zealand.
Statistical analyses reveal surprisingly
little divergence between North Atlantic
and Southern Hemisphere specimens. Co-
efficients of difference for standard meristic
characters are far below accepted levels
for subspecies recognition (all are ^ 0.53),
and regression coefficients for mensural
data are virtually identical. Only maxil-
lary lengths differ significantly at the 95%
level of confidence. It is clear from the
data that North Atlantic and Southern
Hemisphere E. denticulatus do not repre-
sent separate subspecies.
Detailed analyses of the Japanese pop-
ulation could not be undertaken because
of inadequate sampling. Only one speci-
men was available from the area. On the
basis of this fish, the Japanese population
appears closely allied to the rest of the
species. With the exception of eye di-
ameter, standard counts and measm-ements
made on the Japanese morph fall within
the 95% and 99% confidence intervals of
remaining E. denticulatus. Eye diameter
falls outside the 95% confidence interval
but within the 99% confidence interval.
The similarity of E. denficuhiius pop-
ulations, despite the wide rangc> of the
species, suggests ( 1 ) there may be con-
siderable gene flow between populations,
(2) the present distribution may have been
achieved only recently, or (3) evolution
is occurring very slowly. Discovery of a
pelagic juvenile in the mid-North Atlantic
gives credence to the first hypothesis
and proN'ides a mechanism for the dis-
persal of a species with demersal adults
such as E. denticulatus.
Common names. "Castagnera briina" in
Monaco (Bini, 1968).
178 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
W
E
o
o
Q.
a>
m
E
O)
Q.
3
CO
CO
3
T3
T3
C
o
3
CO
V
E
3
CO
3
•a
'>
■D
c
to
o
c
c
o
3
10
b
ii.
Epigonus Systematics • Mayer 179
Epigonus oligolepis sp. nov.
Figure 15
llolotype: One specimen, 90.8 nun SL, taken
from the Straits of Florida bv M/V COMBAT,
Sta. 436: 21 July 1957, 1319 to 1530 hrs.;
24°13'N, 81°42'W; 300 fnis., 10' flat trawl.
USNM 207718.
Parat>pes: One specimen, 126.7 mm SL, M/V
OREGON, Sta. 4731: 27 February 1964;
27°35'N, 92°32'W; 250-300 fms.; 40' flat
trawl. MCZ 48848.
Three specimens, 52.7-72.7 mm SL, Steamer
ALBATROSS, Sta. 2643: 9 April 1886;
25°25'00"N, 79°55'15"W; 211 fms. USNM
109430.
Three specimens, 53.7-84.2 mm SL, M/V
OREGON, Sta. 5043: 26 September 1964;
12°01'N, 6P53.5'W; 210-250 fms.; 40' shrimp
trawl. USNM 207719.
One specimen (cleared and stained), 62.0
mm SL, locality data identical with those of
preceding lot. USNM 207720.
One specimen, 117.1 mm SL, M/V OREGON,
Sta. 3741: 26 August 1962; 29°10'N, 88°01.5'
W; 300-340 fms.; 100' flat trawl. USNM
207721.
Diagnosis. E. oligolepis is distinguished
from all congeners by lateral line scale
counts of 33-36 and the presence of lingual
and endopterygoid teeth.
Description. Meristic values presented in
Table 15; regression data for morphometric
traits presented in Table 16.
Body elongate, moderately compressed;
anterodorsal profile rising gradually from
tip of snout to interorbital region, rising
more steeply and becoming slightly convex
to occiput, thereafter rising gradually to
base of first dorsal fin; body depth 19.8-
24.5% SL; caudal peduncle length 23.9-
27.2% SL.
Head moderate to long, 34.4-43.0% SL;
head height 16.6-18.8% SL; snout pointed;
angle of gape moderate; lower jaw pro-
truding beyond upper jaw. Maxilla reach-
ing %-% eye length; posterior margin
of maxilla rounded, posteriormost point
between midline and ventral margin of
bone. Eye round to slightly oval, 40.1-
43.77o HL; anterodorsal rim of orbit reach-
ing or projecting into dorsal profile; inter-
orbital width 8.5-9.6% SL.
Teeth small, conical; premaxilla edent-
ulous or bearing few teeth on anterior Vi-
-f. of bone; mandibular teeth arranged in
single or double row antericnly, single row
posteriorly; teeth covering anterior half
of bone and occasionally extending along
length of dentaiy. Vomer covered with
irregular tooth patches, teeth extending
posteriorly along midline of palate; pala-
tine teeth arranged in single or multiple
rows anteriorly, single row posteriorly,
covering from half to entire length of bone;
endopterygoid dentigerous; auxiliary tooth
patches occurring between vomer, pala-
tines, and endopterygoids; tongue den-
tigerous, bearing lateral and glossohyal
tooth patches (Fig. lA).
Opercular spine weak, poorly ossified,
ventral to 2-6 membranous spinelets; spine
and spinelets separated by moderate gap;
spinelets occasionally obscured by under-
lying membranes. Preopercular angle rec-
tangular or slightly produced; preopercle,
subopercle and interopercle unserrated.
Gill rakers simple, awl-like.
First dorsal fin VII (10); second dorsal
fin 1,10 (10); anal fin 11,8 (1), 11,9 (9).
Fin spines moderate; DJ 2.7-4.0% SL;
D,I 10.9-12.1% SL; All 10.3-12.2% SL;
PJ 11.0-13.6% SL.
Vertebrae 10 + 15 ( 10 ) , epipleural ribs
7 (4), 8 (1), inserting on vertebrae 1-7
or 1-8 respectively; pleural ribs 7 (10),
inserting on vertebrae 3-9.
Color in alcohol variable with preserva-
tion; specimens frequently abraded reveal-
ing underlying pale yellow or pink-purple
tissue. Recently collected specimens bear
scale pockets mottled with numerous
melanophores; dorsal surfaces of head and
trunk more heavily pigmented; iris black.
Specimens from old collections devoid of
melanin, bearing silver on opercular region,
isthmus, thoracic region, and abdomen to
anus; iris silver. Mouth light, dotted with
brown or black melanophores; l)ranchial
region light in small specimens, darkening
with age.
180 Bulletin Museum of Comparative Zoology, Vol 146, No. 3
r-
o
CM
z
CO
3
CO
E
E
00
o
c>
ui
Q.
>
\-
O
_i
o
Q.
O
to
3
C
O
S.
Uj
in
0)
Epigonus Systematics • Mayer 181
Table 15. Epigonus oligolepis meristic data. X = mean; SD = stan-
dard DEVIATION; n = NUMBER OF SPECIMENS.
X
Range
SD
Pectoral fin rays 17.20 16-18 0.79 10
Gill rakers 30.50 29-31 0.71 10
Lateral line scales 34.70 33-36 1.06 10
Pyloric caeca 8.83 8-10 0.75 6
Table 16. Epigonus oligolepis regression data, b = regression coef-
ficient ± 95% confidence interval; a = Y intercept; n = number of
specimens. All regressions on SL.
b
a
n
HL
0.
36
+
0.
1 1
0.
88
7
Body depth
0.
26
+
0.
02
-2.
45
10
Head height
0.
21
+
0.
06
-2.
56
5
Eye diameter
0.
15
+
0.
03
0.
48
9
Snout length
0.
08
+
0.
03
0.
97
5
Interorbital width
0.
10
+
0.
01
-0.
51
9
Maxillary length
0.
18
+
0,
02
-0.
92
6
Lower jaw length
0.
17
+
0.
02
1.
45
10
Caudal peduncle d
epth
0.
11
+
0.
02
-1.
20
10
Caudal peduncle L
e n g t li
0.
26
+
0.
04
0.
58
9
D2I
0.
13
+
0,
01
-1.
07
6
All
0.
12
+
0.
02
-0.
33
10
P2I
0.
12
+
0.
03
0.
13
9
Description based on 10 specimens 53.7-
126.7 mm SL.
Ontogenetic change. Two juvenile E.
oligolepis (32.0-32.2 mm SL, USNM 207722)
were taken by bottom trawls from
the Gulf of Mexico. These specimens
exhibit many traits characteristic of adult
forms but differ in head shape, meristics.
and dentition. Unlike adults, young E.
oligolepis have smaller eyes (38.2-39.4%
HL) and wider interorbital regions (10.4%
SL). Dorsal fin and gill raker counts are
reduced to VI-I,10 and 26 respectively.
Premaxillary, mandibular, and lingual tooth
patterns are similar to those of mature
individuals, but dentition associated with
182 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
Figure 16. Distribution of E. oligolepis. ■ individual haul of demersal adults; □ individual
haul of demersal juveniles.
the roof of the mouth is strongly reduced.
Vomers and palatines are edentulous or
bear 1-4 teeth; auxiliary tooth patches have
not developed. Endopterygoid teeth are
present but few in number, relatively long,
and medially recurved.
Distribution. E. oligolepis is endemic to
the Gulf of Mexico-Caribbean region (Fig.
16). Specimens have been taken by bot-
tom trawls between 380 and 660 meters.
Remarks. The type specimens of E.
oligolepis exhibit two seemingly disparate
color patterns. One lot, taken in 1886 by
the ALBATROSS, is devoid of melanin but
bears extensive guanine deposits. Remain-
ing fish, all more recently collected, bear
no silver but are dotted with numerous
melanophores. These differences are arti-
facts of preservation.
Specimens collected by early workers
were generally placed directly into ethanol,
while material obtained today is fixed in
10 percent formalin (Hubbs and Lagler,
1958: 16-17). When ethanol is used as a
fixative, it leaches out melanins but does
not affect guanine deposits. Specimens
become pale, but silver pigment is retained.
Formalin has the opposite effect; it
blackens melanophores but destroys gua-
nine crystals. The appearance of preserved
specimens is thus dependent on fixative
composition, concentration, and immersion
time. An alcohol-formalin mixture con-
taining one tablespoon of full strength
formalin per two gallons of 6.5-75 percent
ethanol might be used instead of conven-
tional fixatives to preserve both guanine
and melanin deposits (Myers, personal
communication ) .
Etymology. Oligolepis (Greek), few
scales, from oligos, few, and lepis, scale;
a noun in apposition, refers to the reduced
number of lateral line scales characterizing
the species.
Common names. None.
Epigonvs Systematics • Mayer 183
Figure 17. Epigonus trewavasae, 98.6 mm SL, USNM 207723.
Epigonus trewavasae Poll, 1954
Figure 17
Glossamia pandionis, Lozano (not Goode and
Bean, 1881), 1934: 89; Navarro, 1942: 202;
Navarro et al., 1943: 136, plate XXII, fig. A.
Epigonus trewavasae Poll, 1954: 91, fig. 27
(original description; NOORDENDE III Sta.
52, 06°08'S, 11°30'E, 280-290 m; holotype
examined, IRSN 209).
Epigojitis pandionis, Maurin (not Goode and
Bean, 1881), 1968: 69, fig. 36.
Diagnosis. E. trewavasae is most likely
to be confused with E. robustus, E. leni-
men, E. crassicaudus, and E. pectinifer. It
is distinguished from the first three species
by vertebral counts of 10 + 15 and the
presence of glossohyal and lateral lingual
teeth. The fourth form, E. pectinifer, bears
only glossohyal teeth or a totally eden-
tulous tongue. E. trewavasae may be
further differentiated from E. pectinifer on
the basis of the former's 30-35 awl-like
gill rakers and long, pungent Dol and All
(12.7-16.5% SL, 13.8-16.8% SL respec-
tively). E. trewavasae is unlike remaining
congeners because it bears a pungent, bony
opercular spine, second dorsal fin counts
of 1,9, and pectoral fin counts of 16-18.
Description. Meristic values presented
in Table 17; regression data for morpho-
mctric traits presented in Table 18.
Body elongate; anterodorsal profile flat,
rising without interruption from snout to
base of first dorsal fin; body moderate to
deep, 23.1-27.0% SL; caudal peduncle
length 24.3-27.5% SL.
Head length 33.7-38.1% SL; head height
16.6-18.7%^' SL; snout pointed; angle of
gape small to moderate; lower jaw pro-
truding beyond upper jaw, bearing two
nubs on anterior surface of mandible.
Maxilla reaching slightly less than % eye
length; posterior margin of maxilla narrow,
rounded, or bearing posteriormost point
near midline of bone; short, pungent mus-
tache-like process projecting from postero-
ventral surface of maxillary head. Eye
round, slightly oval in younger specimens,
41.1-49.1%f HL; anterodorsal rim of orbit
reaching profile; interorbital width 8.8-
10.8% SL.
Dentition variable with age (see Onto-
genetic change); teeth conical, small, fre-
quently microscopic, present on premaxiL
lae, mandibles, and vomer; palatines
occasionally edentulous; tongue bearing
lateral and glossohyal tooth patches.
Opercular spine pungent, bony, sur-
mounted by 2-3 horny spinelets; spine and
spinelets separated by large gap; spinelets
often obscured by underlying opercular
membranes. Preopercular angle narrowly
produced, unserrated or bearing serrations
on angle and ventral surface of bone; in-
teropercle and subopercle unserrated or
weakly serrated. Gill rakers simple, awl-
like.
First dorsal fin VII (14); .second dorsal
fin 1,9 (13), 1,10 (1); anal fin 11,9 (14);
DJ moderate, 2.4-3.2% SL; DJ, All, PJ,
184 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
Table 17. Epigonus trewavasae meristic data. X = mean; SD
STANDARD DEVIATION; n =: NUMBER OF SPECIMENS.
X
Range SD
Pectoral fin rays
Gill rakers
Lateral line scales
Pyloric caeca
n
17. 54 16- 18 0. 66 13
33. 15 30- 35 1. 46 13
47. 69 47- 49 0. 75 13
7. 00 6- 8 0. 60 12
Table 18. Epigonus trewavasae regression data, b = regression co-
efficient ±95% confidence interval; a ^ Y intercept; n =: number of
specimens. All regressions on SL.
b
a
n
HL
0. 38
+
0. 03
-2. 03
13
Body depth
0. 29
+
0. 02
-4. 43
12
Head height
0. 19
+
0. 01
-1, 26
12
Eye diameter
0. 17
+
0. 02
-0, 49
13
Snout length
0. 07
+
0. 02
1. 19
13
Interorbital width
0. 09
+
0, 01
1. 41
13
Maxillary length
0. 15
+
0. 02
0. 69
13
Lower jaw length
0. 16
+
0. 01
0. 39
13
Caudal peduncle de
pth
0. 13
+
0. 01
-1. 79
1?
Caudal peduncle len
igth
0, 26
+
0. 02
-0. 08
13
D2 I
0, 15
+
0. 03
-0. 32
12
All
0. 18
+
0. 03
-0. 65
11
P2I
0, 14
+
0. 01
0. 36
13
long, pungent, 12.7-16.5%, 13.(8-16.8%,
13.8-16.27^ SL respectively.
Vertebrae 10 + 15 (12); epipleural ribs
6 (9), 7 (2), inserting on vertebrae 1-6
or 1-7 respectively; pleural ribs 7 (8), 8
(4), inserting on vertebrae 3-9 or 3-10
respectively.
Color variable with presei'vation; speci-
mens abraded, revealing underlying yel-
low to yellow-pink tissue; fin membranes
dark; scale pockets covered with dense
brown or black melanophores; dorsal sur-
face of trunk more heavily pigmented than
ventral; opercles brown, black, or slate
gray; guanine deposits occurring occasion-
ally on opercular region and from isthmus
to bases of paired fins; iris black with sil-
ver highlights; mouth light; branchial re-
Epigonus Systematics • Mayer 185
JO' SO"
Figure 18. Distributions of E. trewavasae and E. pectinifer. Large map shows localities in the Atlantic; insert
shows localities off Japan. E. trewavasae: ^ individual haul of adults; Q individual haul of juveniles; cross-
hatching indicates areas of capture cited in the literature. £. pectinifer: ■ individual haul of adults; □ individ-
ual haul of juveniles; A report from the literature.
gion light in smiill specimens, becoming
l)lack with age.
Description based on 13 specimens 70.9-
153.9 mm SL.
Ontogenetic change. The most striking
ontogenetic changes in E. trewavasae are
associated with the development of adult
tooth patterns. Large specimens bear ir-
regular double or triple rows of premaxil-
lary and mandibular teeth that taper to a
single row posteriorly. Vomers are covered
with minute conical teeth, while palatines
are either edentulous or bear single to
double rows of teeth.
Dentition patterns are simple in small
specimens but become more complex as
teeth are added during growth. A 29.8-
mm juvenile lacks both premaxillary and
mandibular teeth. By 70-75 mm SL teeth
are present in single rows on the jaws, and
by 145 mm SL adult tooth patterns pre-
vail. As premaxillary tooth patches widen,
they extend posteriorly and eventually
cover the first half of the bone. Analogous
expansion occurs in vomerine tooth
patches.
Distribution. E. tretcavasae is known
from equatorial west Africa, northwest
Africa, and the western Mediterranean
(Fig. 18). It has been taken by bottom
trawls between 200 and 600 meters.
Geographic variation. Statistical com-
parisons of African and Mediterranean E.
trewavasae were not made because of
small sample size. As additional material
is collected, the following intraspecific
differences should be examined:
( 1 ) vomerine and palatine teeth more
strongly developed in Mediterra-
nean forms;
(2) chin nubs more strongly developed
in African forms;
(3) preopercular serrations more
strongly developed in Mediterra-
nean forms.
Although the significance of thc\se features
186 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
is unknown, they suggest that African and
Mediterranean forms may represent sepa-
rate subspecies.
Taxonomic notes. Pomatomichthys con-
stonciae GigHoh, 1880 may be a synonym
of E. trewavasae Poll, 1954. See E. tele-
scopus: Taxonomic notes, for a discussion
of this possibility.
Remarks. Dieuzeide (1950: 104-105)
reported that specimens designated as
Glossamia pandionis hy hozano (1934) and
NavaiTO et al. (1943) were actually mis-
identified E. denticulatus. This is incor-
rect. Lozano's report is based on a single
specimen (131 mm total length) taken
from the Catillian coast. Among the char-
acters cited for this fish are dorsal fin
counts of VII-1,9, pectoral counts of 16,
and an All subequal to the eye diameter
(p. 89). All of these are characters diag-
nostic of E. trewavasae. E. denticulatus
bears 10 rays in the second dorsal fin, 18-
20 pectoral rays, and an All equalling half
the eye diameter.
Navarro et al.'s specimens also appear
to be E. trewavasae. Altliough no descrip-
tion is provided, the account includes a
photograph (plate XXII, fig. A) that shows
the fish have deep bodies, pungent oper-
cular spines, and long D2I, All, and P2rs.
All of these features are characteristic of
E. trewavasae.
More recently, Maurin (1968) mistook
E. trewavasae for E. pandionis. Propor-
tional measurements of body depth, head
height. All, and P-I made on Maurin's
figure 36 (p. 69) fall within ranges char-
acteristic of E. treioavasae; however, pub-
lished gill raker counts of 28-30 (p. 70)
are lower than expected.
Common names. None.
Epigonus pectin! fer sp. nov.
Figure 19
Ilolotype: A 114.3-mm SL specimen taken from
the Caribbean west of Grenada by M/V
OREGON, Sta. 5043: 26 September 1964,
12°01'N, 61°53.5'W, 210-250 fms., 40' shrimp
trawl. USNM 207725.
Paratypes: One specimen, 97.4 mm SL, 16
September 1964, Suruga Bay, commercial trawl.
ABE 64-2085.
One specimen, 100.6 mm SL, 14-31 October
1964, Suruga Bay, commercial trawl. ABE
64-2245.
One specimen, 99.8 mm SL, 14-31 October
1964, Suruga Bay, commercial trawl. ABE
64-2248.
Two specimens, 95.2-117.1 mm SL, station
data identical with those of holotype. MCZ
48850.
One specimen (cleared and stained), 108.1
mm SL, station data identical with those of
holotype. MCZ 48851.
One specimen, 94.8 mm SL, R/V PILLS-
BURY, Sta. P-582: 23 May 1967; 21°10'N,
86°18'W; 250-155 fms.; 10' otter trawl. UMML
30378.
One specimen, 111.2 mm SL, M/V OREGON,
Sta. 4405: 27 September 1963; 11°53'N,
69°28"W; 215 fms.; 40' flat trawl. USNM
207726.
Ten specimens, 101.8-120.6 mm SL, station
data identical with those of holotype. USNM
207727.
Nine specimens 81.5-118.9 mm SL, station
data identical with those of holotype. USNM
207728.
Two specimens (cleared and stained), 94.8-
98 mm SL, station data identical with those
of holotype. USNM 207729.
Epigonus rohiistiis, Matsubara (not Barnard,
1927), 1936: 121, fig. IB; Kamohara, 1952:
37.
Diagnosis. E. pectinifer is characterized
by comb-like gill rakers on the lower half
of the first gill arch. This feature, together
with glossohyal dentition (present in most
specimens) and vertebral counts of 10 +
15, differentiate E. pectinifer from E. ro-
htistus, E. lenimen, and E. crassicaudus.
E. pectinifer most closely resembles E. tre-
ioavasae but is distinguished by less exten-
sive lingual dentition, fewer gill rakers
(26-30), and shorter DJ and All (11.2-
12.7% SL and 11.9-14.0% SL respectively).
E. pectinifer may be separated from re-
maining congeners by its pungent, bony
opercular spine, second dorsal fin counts
of 1,9, and pectoral fin counts of 15-18.
Description. Meristic values presented
in Table 19; regression data for morpho-
metric traits presented in Table 20.
Epigonus Systematics • Mayer 187
CM
o
CM
3
E
E
UJ
Q.
>-
!^
<
Q.
O
V>
Q.
in
C
O
o>
188 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
Table 19. Epigonus pectinifer meristic data. X = mean; SD = stan-
dard DEVIATION; n = NUMBER OF SPECIMENS.
Range SD
Pectoral fin rays 16.03 15-18 0.57 29
Gill rakers 27.59 26-30 0.98 29
Lateral line scales 48.14 47-49 0.58 29
Pyloric caeca 6, 10 5- 7 0. 41 29
Table 20. Epigonus pectinifer regression data, b = regression coef-
ficient ± 95% confidence interval; a = Y intercept; n = number of
specimens. All regressions on SL.
b
a
n
HL
0.
35
+
0.
05
-2.
21
27
Body depth
0.
28
+
0.
03
-4.
95
28
Head height
0.
18
+
0.
03
-2.
83
19
Eye diameter
0,
17
+
0.
03
-3.
18
28
Snout length
0.
09
+
0.
03
0.
03
26
Interorbital width
0.
10
+
0.
03
-1,
84
28
Maxillary length
0.
17
+
0.
04
-2.
10
26
Lower jaw length
0.
15
+
0.
03
0.
55
28
Caudal peduncle de
pth
0.
84
+
0,
03
1.
88
28
C audal peduncle length
0.
27
+
0.
04
-0.
22
28
D2I
0.
11
+
0.
02
0.
62
24
All
0.
12
+
0.
02
0.
32
24
P2I
0.
11
+
0.
02
1.
30
28
Body elongate; anterodor.sal profile flat
or slighdy convex, rising withont interrup-
tion from snout to base of first dorsal fin;
body depth 21.1-24.6% SL; caudal pedun-
cle narrow, length 2.5.1-28.7% SL.
Head short to moderate, 31.3-35.7% SL,
shallow, 14.2-16.9% SL; snout wide,
pointed; angle of gape small; lower jaw
proti-uding slightly beyond upper jaw;
nubs at anterior end of mandible paired,
barely discernible, or absent. Maxilla
reacliing %-y2 eye length, posterior margin
narrow, rounded, or bearing posteriormost
point near midline of bone; short, pun-
gent, mustache-like process projecting
from posteroventral surface of maxillary
head. Eye round or slightly oval, 38.7-
45.4% HL; anteiodorsal rim of orbit not
Epigonus Systematics • Maijer 189
reaching profile; intcrorl)ital widtli 7.7-
9.4% SL.
Teeth small, conical; premaxilla edentii-
lons or bearing teeth anteriorly; when
present, teeth 1-15, arranged in single
row. Mandibular teeth covering all or part
oi dentary, arranged in single row. Vomer-
ine teeth strong, arranged in tightly packed
()\'al patch. Palatines edentulous or bear-
ing teeth anteriorly; when present, teeth
1-6, arranged in single row; tongue with
glossohyal teeth, rarely edentulous.
Opercular spine pungent, bony, ventral
to 2-3 horny spinelets; spine and spinelets
separated by large gap; spinelets occasion-
ally obscured by underlying membranes.
Preopercular angle narrowly produced,
serrated; subopercle and interopercle un-
serrated or weakly serrated. Gill rakers
pectinate, bearing nub-like projections
proximally along mesial surfaces (Fig.
IC); pectinate structure variable in extent,
most prominent on ventral portions of gill
arch.
First dorsal fin VII (28); second dorsal
fin 1,9 (29); anal fin 11,9 (29); D,I
short, 1.6-2.8% SL; D,I moderate, 11.2-
12.7% SL; All, PJ, 11.9-14.0% SL.
Vertebrae 10 + 15 (29); epipleural ribs
6 (17), 7 (13), inserting on vertebrae 1-6
or 1-7 respectively; pleural ribs 8 (29),
inserting on vertebrae 3-10.
Color in alcohol brown-black; fin mem-
branes black; scale pockets covered with
densely packed melanophores; skin often
abraded, revealing underlying yellow-pink
tissue; iris black; branchial region white to
dark gray; mouth light.
Description based on 30 specimens 81.5-
120.6 mm SL.
OntO(!,enetic change. A 33.8-mm E. pec-
tinifer was taken by bottom trawl in the
Gulf of Mexico (USNM 207731). The
specimen appears similar to adults and pro-
vides little evidence of ontogenetic change.
The major difference is the presence of six
rather than seven first dorsal fin spines.
Distribution. E. pectinifer is known
from the Caribbean Sea, Gulf of Mexico,
and eastern coast of Japan (Fig. 18).
Specimens were taken between 280 and
550 meters.
GeograpJiic variation. Definitive com-
parisons of Japanese and American E. pec-
tinifer were not undertaken, because only
three oriental specimens were available for
study. The latter forms were, however, in-
dividually compared with Amcnican fish.
The analyses revealed virtually no differ-
ences between the populations aside from
a slight tendency toward broader caudal
peduncles and shorter maxillae and man-
dibles by the Japanese specimens.
Remarks. A teratological specimen of
E. pectinifer was taken from the Yucatan
Channel (109 mm SL, UMML 30379). The
fish was captured at depths characteristic
of E. pectinifer and bears diagnostic traits
such as 27 gill rakers ( many are pectinate ) ,
VII + I dorsal fin spines, 16 pectoral fin
rays, and 10 + 15 vertebrae. The tongue
is edentulous. Unlike the condition in
typical forms, opercles are not fully ossified
and lack spines and spinelets. Similarly,
the lateral line is incomplete on the right
side and bears only 43 pored scales on the
left. Other differences include enlarged
teeth and chin nubs, 10 rather than 9 dor-
sal rays, and 8 rather than 5-7 pyloric
caeca.
The aberrant specimen was not consid-
ered in preparing the species description.
Etymology. Pectinifer (Latin), comb-
bearer, from pecten, comb, and ferare, to
bear; a noun in apposition, refers to the
comb-like gill rakers characterizing this
species.
Common names. None.
Epigonus robustus (Barnard, 1927)
Figure 20
Epigonus macrops Gilchrist and von Bonde, 1924:
14, plate I, fig. 3 (oiij^inal description; S..S.
PICKLE Sta. 344, 30°12'00"S, 14°25'()()"E,
510 fms.; Sta. 347, 31°58'00"S, 16°00'00"E,
670 fms.; syntype examined, RUSI 669; name
suppressed, junior homonym of Oxi/odot} macrops
Brauer, 1906); Barnard, 1927:' 523; Smith,
1961: 377, fig. 2.
190 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
Figure 20. Epigonus robustus, 154.6 mm SL, LACM 11449-7.
Parahynnodiis robustus Barnard, 1927: 525, plate
XXII, fig. 4 (original description; off Cape
Point, 460 fms.; holotype in poor condition, not
examined, SAM 13080).
Hymwdus robustus Smith, 1949b: 210, fig. 495.
Diagnosis. E. robustus sti-ongly resem-
bles E. pectinifer, E. trewavasae, and E.
lenimen. It may be distinguished from
the former two species by the absence of
hngual teeth. In addition, mihke E. pec-
tinifer, it has awl-Hke gill rakers. E. ro-
hu^us differs from E. lenimen by having
a narrow interorbital region (6.5-8.2%
SL), short DJ (10.0-12.6% SL) and short
All (9.2-13.3% SL). E. robustus may be
distinguished from E. crassicaudus by the
former's short head (28.0-34.0% SL) and
shallow body (20.3-24.6% SL). It differs
from remaining congeners by bearing a
pungent, bony opercular spine, vertebral
count of 11 + 14, and nine rays in the
second dorsal fin.
Description. Meristic values presented
in Table 21; regression data for morpho-
metric traits presented in Table 22.
Body elongate, moderately compressed;
anterodorsal profile weakly convex, rising
without interruption from tip of snout to
base of first dorsal fin; body depth 20.3-
24.6% SL; caudal peduncle moderate to
long, 25.3-30.7% SL.
Head short, shallow, length 28.0-34.0%
SL, height 14.8-16.3% SL; snout short,
pointed; angle of gape moderate to large;
lower jaw protruding beyond upper jaw,
bearing two nubs of variable prominence
on anterior surface of mandible. Maxilla
reaching ¥3-^/4 eye length; posterior margin
of maxilla narrow, rounded or bearing
posteriormost point near midline of bone;
small, weak mustache-like process project-
ing from j)osteroventral surface of maxil-
lary head. Eye round to oval, small, 37.4-
42.4% HL; anterodorsal rim of orbit not
reaching dorsal profile; interorbital region
narrow, 6.5-8.2% SL.
Teeth small, conical; premaxilla edentu-
lous or bearing single row of teeth on an-
terior half of bone; mandibular dentition
covering all or part of dentary, arranged
in double row anteriorly, tapering to single
row posteriorly; vomer bearing 1-6 irregu-
lar rows of teeth; palatines edentulous or
bearing teeth on anterior half of bone;
tongue edentulous.
Opercular spine pungent, bony, ventral
to 2-3 membranous or horny spinelets;
spine separated from spinelets by wide
gap; spinelets often obscured by underly-
ing membranes. Preopercular angle not
produced, serrations on posterior and/ or
ventral surfaces of bone rarely absent; sub-
opercle and interopercle serrated. Gill
rakers simple, awl-like.
First dorsal fin VI (1), VII (27), VIII
(1); second dorsiil fin I, 9 (28), II, 8 (1);
anal fin II, 9 (29). DJ short, 1.4-2.5%
SL; DJ, All, PJ moderate to long, 10.0-
12.6%, 9.2-13.3%, 11.7-15.3% SL respec-
tively.
Vertebrae 11 + 14 (29); epipleural ribs
Epigonus Systematics • Mayer 191
Table 21. Epigonus robustus meristic data. X = mean; SD = stan-
dard DEVIATION; n = NUMBER OF SPECrMENS.
X Range SD n
Pectoral fin rays 16.79 16-18 0.55 29
Gill rakers 31.68 30-33 0.93 29
Lateral line scales 48.76 47-50 0.91 29
Pyloric caeca 6.36 5- 8 0.78 28
Table 22. Epigonus robustus regression data, b = regression coef-
ficient ± 95% confidence interval; a = Y intercept; n = number of
specimens. All regressions on SL.
b
a
n
HL
0.
28
+
0.
02
5,
35
28
Body depth
0.
28
+
0.
02
-7.
80
28
Head height
0,
17
+
0.
02
- 1.
65
20
Eye diameter
0.
11
+
0.
01
44
28
Snout length
0.
06
+
0.
02
80
23
Interorbital widtl:
0.
09
+
0.
01
- 1.
97
28
Maxillary length
0.
12
+
0.
01
14
24
Lower jaw length
0,
14
+
0.
01
27
27
Caudal peduncle dep
ith
0.
13
+
0.
01
- 1.
69
28
Caudal peduncle len
gth
0.
25
+
0.
02
5.
32
28
D2I
0.
08
+
o„
02
5.
49
2 1
All
0.
07
+
0,
02
8„
18
15
P2I
0.
09
+
0.
02
6.
74
24
6 (2), 7 (8), inserting on vertebrae 1-6 brown or black niclanopliores; brancliial
or 1-7 respectively; pleural ribs 9 (29), in- region black. Body very oily; body cavity
serting on vertebrae 3-11. filled with rust brown fat globules; viscera
Color variable with preservation, pale and swinibladder often completely envel-
yellow to rust brown; scale pockets out- oped in fat.
lined by small black or brown melano- Description based on 29 .specimens
phores; opercular region tinged with 121.1-198.0 mm SL.
black; iris black; mouth light, mottled with Distribution. Most specimens of E. ro-
192 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
EriGONus Systematics • Mayer 193
Figure 22. Epigonus lenimen, 139.0 mm SL, UZM P45165.
biistus liave been taken by bottom trawLs
between 800 and 1225 meters off south-
eastern South Ameriea, South Africa, and
AustraHa (sec Fig. 21). One specimen
(ISH 430/71) was taken by a deep pelagic
trawl.
Geographic variation. No investigation
was undertaken because insufficient ma-
terial was available from South Africa and
Australia.
Taxonomic notes. Epigonus macrops
Gilchrist and von Bonde, 1924 was des-
cribed from two syntypes; the larger was
19(S mm (SL?). These specimens, together
with manv others collected bv the Fish-
cries and Marine Biological Survey, were
lost while being transferred to the South
African Museum. A portion of the ma-
terial was subsequently rediscovered at
Rhodes University, Grahamstown. From
the contents Smith (1961: 378) described
a specimen that he believed to be "Gil-
christ and \'on Bonde's type of macrops
from 600 fathoms off St. Helena Bay." This
fish was re-examined during the present
study.
Smith's specimen measures 162.2 mm SL
and thus cannot be the larger syntype;
however, it conforms to the descriptions
and proportions supplied by Gilchrist and
von Bonde and probably represents the
smaller type for which no length was pub-
lished.
An unusual aspect of the syntypes is that
the locations at which they were captured
will never be precisely known. The speci-
mens were taken at different stations. Al-
though these are recorded in both the orig-
inal description of E. macrops and in the
1921 report of the Fisheries and Marine
Biological Survey (Gilchrist, 1922), neither
account specifies which data are associated
with which syntype.
Common names. None.
Epigonus lenimen (Whitley, 1935)
Figure 22
Scepterias lenimen WHiitley (in part), 1935: 230
(original description; Great Australian Biglit:
south from Eucla, 350—450 fnis.; holot\-pe
examined, AM E3368); Whitley, 1940: 420,
fig. 33; Wliitley (in part), 1968: 56.
Epigonus lenimen Scott, 1962: 191, 1 fig.
Diagnosis. E. lenitnen is distinguished
from E. robustus and E. crassicaudus by
its broad interorbital region (8.7-10.2%
SL), long DJ (14.9-18.7% SL), and large
eyes (40.0-51.1% HL). It is further dif-
ferentiated from E. crassicaudus by shorter
head lengths (32.7-36.67^ SL) and .shal-
lower head heights (16.2-18.8% SL). E.
lenimen lacks lingual teeth but has 11 + 14
vertebrae and thus may be distinguished
from E. trewavasae and E. pectinifer. Un-
like remaining congeners, E. lenimen bears
a pungent, bony opercular spine, nine
second dorsal fin rays, and 16-18 pectoral
fin rays.
194 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
Table 23. Epigonus lenimen meristic data. X = mean; SD = standard
deviation; n = nxtmber of specimens.
X
Range
SD
Pectoral fin rays 16.96 16-18 0.58 28
Gill rakers 30.29 28-34 1.27 24
Lateral line scales 48.12 47-50 0.91 26
Pyloric caeca 7.33 7- 9 0.56 24
Table 24. Epigonus lenimen regression data, b = regression coeffi-
cient ± 95% confidence interval; a = Y intercept; n = nltmber of
specimens. All regressions on SL.
b
a
n
HL
0.
35
+
0,
01
-0.
12
29
Body depth
0.
28
+
0.
02
-3.
32
27
Head height
0,
19
+
0.
01
-1.
08
27
Eye diameter
0.
18
+
0.
01
-1,
60
27
Snout length
0.
08
+
0.
01
0.
31
27
Interorbital width
0.
10
+
0,
01
-0.
59
26
Maxillary length
0.
16
+
0.
01
0.
08
27
Lower jaw length
0.
16
+
0.
01
0.
55
28
Caudal peduncle dep
th
0.
11
+
0.
01
-0.
14
28
Caudal peduncle len
gth
0.
24
+
0.
02
2.
26
26
D2I
0.
17
+
0.
02
-0.
79
18
All
0.
21
+
0.
02
-2.
36
22
P2I
0.
19
+
0.
01
-2,
13
28
Description. MerLstic values presented in
Table 23; regression data for morphometric
traits presented in Table 24.
Body elongate; anterodorsal profile flat
or weakly concave, rising without inter-
ruption to first dorsal fin, more steeply
inclined behind occiput in large specimens;
body moderate to deep, 21.5-27.5% SL;
caudal peduncle moderate to long, 23.6-
29.3% SL. Head length 32.7-36.6% SL;
head height 16.2-18.8% SL; snout moder-
ately pointed; angle of gape moderate,
variable with age; lower jaw protruding
slightly or not at all; no prominent nubs on
anterior surface of mandible. Maxilla
reaching Vs-V2 eye length; posterior margin
Epigonus Systematics • Mayer 195
Table 25. Comparison of E. LENiMEy paratypes with specimens of E. lknimen and E.
DENTICULATVS. PaRATYPE MERISTICS REPORTED AS VALUE, FOLLOWED IN PARENTHESES HY NUM-
BER OF SPECLMENS EXHIBITING THAT VALUE. RATIOS ARE EXPRESSED AS PERCENTAGES.
E . 1 e n i m e n
E_. 1 e n i m e n paratypes E^. denticulatus
Dorsal fin rays 8-9 9(1), 10(11) 10
Pectoral fin rays 16—18 19(6), 20(6) 18—20
Vertebrae 11+14 10+15 10+15
10(1), 11(3) , „ , ,
Pyloric caeca 7 — 9 12(7) —
BH/SL 21.5—27.5 18.4 — 21.7 15.8—23.6
D2 I/SL 14.9—18.7 6.0—7.6 5.3—8.0
AII/SL 13.0 — 20.8 6.2—7.1 6.0—8.2
P2 I/SL 12.5 — 18.7 8.5—9.9 7.9—10.0
of maxilla narrow, rounded, or bearing First dorsal fin VII (29); second dorsal
posteriormost point near midline of bone; fin 1,8 (1), 1,9 (28); anal fin 11,8 (2), 11,9
weak mustache-like process projecting from (26); DJ moderate, 2.0-4. 17^ SL; DJ,
posteroventral surface of maxillary head, All long, 14.9-18.7%, 13.0-20.8% SL re-
process occasionally absent. Eye large, spectively; PJ moderate to long, 12.3-
oval, 40.0-51.1% HL; anterodorsal rim of 18.7% SL.
orbit reaching dorsal profile; interorbital Vertebrae 11 + 14 (29); epipleural ribs
width 8.7-10.2% SL. 6 (6), 7 (12), 8 (2), inserting on vertebrae
Teeth small, conical; premaxilla eden- 1-6, 1-7, or 1-8 respectively; pleural ribs
tulous or bearing single row of teeth oc- 9 (28), inserting on vertebrae 3-11.
cupying anterior half of bone. Mandible Color in alcohol variable; skin often
edentulous or bearing single row of teeth abraded, revealing underlying pale pink-
occupying up to % of dentary; tooth row yellow tissue; fin membranes and scale
occasionally double near symphysis. Vo- pockets mottled with numerous black
mer edentulous or bearing up to seventeen melanophores; head, opercular region, and
teeth arranged in diamond-shaped patch fin bases deep rust brown. Guanine de-
or in 1-3 irregular rows; palatines edentvi- posits variable, occurring on ventral por-
lous or bearing 1-2 teeth anteriorly; tongue tions of opercular region, isthmus, pectoral
edentulous. and pelvic fin bases, and abdomen to anus;
Opercular spine pungent, bony, ventral silver chromatophores on dorsal, anal, pee-
to 1-5 (usually 2) membranous or horny toral, or pelvic fin rays; iris black with
spinelets; spine and spinelets separated by silver highlights; moutli light, dotted with
wide gap; spinelets frequently obscured by melanophores; branchial region light in
underlying membranes. Preopercular angle small specimcMis, blackening with age.
narrowly produced, occasionally serrated; Description based on 32 specimens 40.0-
subopercle and interopercle unserrated or 147.8 mm SL.
weakly serrated. Gill rakers simple, awl- Distrihution. E. lenimen is known from
like. three localities (Fig. 21). The liolotype
196 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
0 I 2 3 cm
Figure 23. Epigonus crassicaudus, 259.0 mm SL, MCZ 48855.
was taken south of Austi-alia between 622
and 823 meters. Remaining specimens
were taken off New Zealand between 530
and 660 meters.
Taxonomic notes. Although G. P. Whit-
ley was the first to describe E. lenimen
(1935), inaccuracies in his papers have
produced several problems. The most
serious involve the type series and type
locality of the species.
The original description of E. lenimen
designates a holotype and nine paratypes.
A figure of the new form was not included
but was published in a subsequent paper
(Whidcy, 1940: fig. 33). Both the de-
scription and the illustration were based
exclusively on the holotype. Whitley re-
alized that the paratypes were different
from the holotype but considered them
to be poorly preserved specimens ( Whitley,
1935: 320).
A re-examination of the type series re-
veals that the paratypes are not conspecific
with the holotype. They are, instead, mem-
bers of E. denticulattis. As is shown in
Table 25, counts and measurements from
the paratypes always fall within ranges
characteristics of E. denticulatus. Pectoral
fin counts, vertebral number, pyloric
caecum counts, and fin spine lengths are
particularly noteworthy in this respect.
The paratypes further resemble E. denti-
culatus by bearing dentigerous palatines
and weak opercular armor, E. lenimen, on
the other hand, is characterized by eden-
tulous palatines and pungent opercular
spines.
Confusion over the type locality stems
from Whitley's 1940 paper. The locality
is cited as "from 190-320 fathoms, S.W.
from Eucla, Great Australian Bight [p.
420] ." This contradicts the data presented
in the original description: "Great Aus-
ti-alian Bight; south from Eucla, 350 to 450
fathoms [p. 231]." The 1940 citation is
extremely similar to station data listed for
paratypes AM E3581-3582 in 1935 ("Great
Australian Bight; SW from Eucla, 190-320
fathoms. 126° 451/2'E long. [p. 231]"). In
the absence of other information, it must be
concluded that erroneous locality data were
inserted in the 1940 publication through an
editorial oversight.
The most recent taxonomic questions
arise from Whitley's check list of New
Zealand fishes ( 1968 ) . This work includes
two incorrect citations in the synonymy
of E. lenimen. The first is based on a fish
taken off the Chatham Islands and tenta-
tively identified as Hynnodiis atherinoides
(Moreland, 1957). This specimen was later
re-identified as Grahamichthijs radiatus
(Moreland, personal communication). The
second misidentified .specimen is a "Big-
eyed Cardinal Fish" captured off Cape
Palliser, New Zealand (Anonymous, 1961).
This fish is actually a specimen of E.
telescopus and is presently in the collec-
Ei'ic.oNus Systematics • Mayer 197
tions of the Dominion Museum (DM 3072,
examined ) .
Common names. None.
Epigonus crassicaudus de Buen, 1959
Figure 23
Epiguinis crcis.'iicattdii.s de Buen, 1959: 196
(original description; preabysnial zone off
Valparaiso, Chile; holotype not examined, EBM
10.183).
Diagnosis. E. crassicaudus is strongly
compressed. It reaches 260-270 mm SL
and is the second largest species in the
genus. E. crassicaudus may be distin-
guished from E. trewavasae, E. pectinifer,
E. rohustus, and E. lenimen by its deep
head (18.9-21.2% SL) and deep body
(24.3-32.0% SL). It differs from remain-
ing congeners by bearing 9 rays in the
second dorsal fin and 6-7 pyloric caeca.
Description. Meristic values presented
in Table 26; regression data for morpho-
metric traits presented in Table 27.
Body elongate, compressed; anterodorsal
profile rising from tip of snout to occiput,
becoming moderately convex from occiput
to base of first dorsal fin. Body deep, 24.3-
32.07f SL; caudal peduncle broad, moderate
to short, 21.6-26.4% SL.
Head long, deep, postorbital portion
greatly expanded, length 36.8-41.9% SL;
height 18.9-21.27r SL; snout moderately
pointed in small specimens, blunt in adults;
angle of gape moderate to small; mandible
long, strongly protuberant, young bearing
two weak nubs on anterior surface of lower
jaw. Maxilla reaching %-% eye length;
posterior margin of maxilla broad, rounded
or bearing posteriormost point between
midline and ventral surface of bone. Eye
round, small, 34.2-39.6% IlL; surrounded
by numerous small scale pockets; antero-
dorsal rim of orbit reaching dorsal profile^;
interorbital region narrow, 6.2-8.5% SL.
Teeth small, conical, occasionally villi-
form, larger in small specimens; premax-
illary teeth arranged in irregular single
or double rows tapering to single row
posteriorly and covering from % to entire
length of bone; mandibular teeth arranged
in multiple rows, tapering to single row
posteriorly, covering from V2 to entire
IcMigtli of dentary; vomer edentulous or
bearing up to six irregular rows of minute
teeth; palatines edentulous or bearing 1-3
teeth anteriorly; tongue edentulous.
Opercular spine pungent, bony, ventral
to 3-5 flat, horny spinelets; spine separated
from spinelets by narrow gap; spinelets
often obscured by underlying membranes.
Preopercular angle slightly produced, pos-
terior and/ or venti'al surfaces serrated;
subopercles and interopercles serrated.
Gill rakers awl-like, short; gill filaments
long.
First dorsal fin VII (22); second dorsal
fin 1,9 (20), 1,10 (2); anal fin II.8 (1),
11,9 (21); DJ 2.0-3.6% SL; DJ 9.-8-13.2%
SL; All 10.3-14.0% SL; PJ 13.0-15.5%
SL.
Vertebrae 11 + 14 (25); epipleural ribs
6 (2), 7 (16), inserting on vertebrae 1-6
or 1-7 respectively; pleural ribs 9 (25),
inserting on vertebrae 3-11.
Color in alcohol variable with preserva-
tion; skin frequently abraded, exposing
underlying pink tissue and orange-rust fat
deposits; skin exti'emely oily; fin membranes
black; scale pockets mottled with numerous
black melanophores; dorsal portion of body
darker than ventral; forehead, snout, an-
terior half of mandible, and circumorbital
area heavily invested with black pigment;
opercles black or slate gray. Guanine de-
posits occasionally on opercles, isthmus,
pectoral and pelvic fin bases, and al)do-
men to anal fin; iris variable — black, siher,
or black with silver highlights; mouth and
branchial region light, darkening with age.
Description based on 27 specimens 80.3-
262.5 mm SL.
Ontogenetic change. Two juvenile E.
crassicaudus (12.2 mm SL, MCZ 48857,
and 21.7 mm SL, MCZ 48858) were taken
off the Chilean coast by midwater trawl.
Although these forms bear characteristics
diagnostic of the species, they differ con-
198 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
Table 26. Epigonus crassicaudus meristic data. X = mean; SD
STANfDARD DEVIATION; n = NUMBER OF SPECIMENS.
X
Range
SD
Pectoral fin rays 18.05 17-19 0.58 22
Gill rakers 32.27 31-34 0.70 22
Lateral line scales 47.86 46-49 0.85 21
Pyloric caeca 6.87 6- 7 0.35 15
Table 27. Epigonus crassicaudus regression data, b = regression
COEFFICIENT ± 95% CONFIDENCE INTERVAL; a = Y INTERCEPT; n = NUMBER
OF SPECIMENS. AlL REGRESSIONS ON SL.
b
a
n
HL
0.
39
+
0.
02
0.
47
20
Body depth
0.
30
+
0.
04
-3.
75
21
Head height
0.
21
+
0.
02
-1.
62
18
Eye diameter
0.
14
+
0.
01
0.
74
21
Snout length
0.
08
+
0.
01
0.
08
19
Interorbital width
NONLINEAR
Maxillary length
0,
17
+
0.
01
-0.
49
20
Lower jaw length
0.
20
+
0,
01
-1,
17
22
Caudal peduncle dep
th
0,
12
+
0.
01
- 0,
65
20
Caudal peduncle len
gth
0.
22
+
0,
03
4,
32
20
D2 I
0.
10
+
0.
02
3.
34
12
All
0.
11
+
0.
02
2.
63
17
P2I
0.
14
+
0.
01
0.
05
16
sideral)ly in appearance and habit from
adnlts.
Most .striking is the juvenile pigment
pattern. Pelagic specimens are basically
pale yellow with large, brown patches
covering most of the caudal peduncle.
Caudal peduncle rings, like those found on
E. pandionis young, are absent, although
myotomes are outlined by thin brown
bands. Brown pigment extends anteriorly
as a band from the caudal peduncle to the
frontal region of the head. A poorly de-
fined black stripe extends across the snout
to the anterior rim of the orbit. In general,
juvenile E. crassicaudus resemble E. teles-
copiis young figured by Koefoed (1952:
plate IIA).
The midwater capture of E. crassicaudus
Epigonus Systematics • Mayer 199
juveniles suggests that tlie life cycle of tlK>
specit^s includes a pelagic stage. Unfortu-
nately, the data available are not sufficient
to determine the duration of this stage.
Distribution. E. crassicaudus is endemic
to the waters off central Chile (Fig. 21).
Adults have been captured by bottom
trawls made between 200 and 400 meters;
juveniles were taken by midwater trawls
fishing from 200 to 270 meters.
Common mimes. None.
Species Incertae Sedis
Micwichtlnjs coccoi Riippell, 1852: 1 (original
description; "Mare siculum"; holotype not ex-
amined, SMF 1069).
The original description of M. coccoi
provides only a superficial account of the
holotype. Subsequent papers either para-
phrase Riippell's work {e.g., Canestrini,
1860; Doderlein, 1889) or are based on
material not compared to the holotype
(i.e., Facciola, 1900; Caporaicco, 1926;
Gonzales, 1946). It is questionable whether
the latter specimens are conspecific with
the holotype.
Most recent revisers (e.g., Schultz, 1940;
Norman, 1957) have synonymized Micro-
ichthys with Apogon; however, the data
are inconclusive and also suggest an affinity
with Epigonus (Eraser, 1972: 5). A re-
examination of the holotype must be under-
taken to clarify the status of M. coccoi.
A second species of Microichthys — M.
sonzoi Sparta, 1950 — does not appear to be
an Epigonus on the basis of vertebral and
dorsal fin counts. The only known speci-
men of this species has been lost (Torton-
ese, personal communication).
ACKNOWLEDGMENTS
This work would not have been possible
without the assistance and support of
niunerous people. I wish to thank the
following scientists and institutions for
material used in this study: J. R. Paxton
and D. Hoese, Australian Museum; A. W.
Wheeler and G. Palmer, British Museum
(Natural History); J. Randall, Bemice P.
Bishop Museum; W. Eschmeyer, California
Academy of Sciences; E. Bertclsen, Carls-
bergfondets; J. Moreland, Dominion Mu-
seum; L. P. Woods, Eield Museum of
Natural History; P. Struhsaker, National
Marine Fisheries Service, Honolulu; R.
Raymond, Instituto de Fomento Pesquero;
X. Missonne, Institut Royal des Sciences
Naturelles de Belgique; G. Krefft, Institut
fiir Seefischerei; M. M. Smith, J. L. B.
Smith Institute of Ichthyology; I. Naka-
mura, Kyoto University; R. J. Lavenberg,
Los Angeles County Museum of Natural
History; M. Bauchot, Museum National
d'Histoire Naturelle; M. Poll, Musee Royal
de I'Afrique Centrale; E. A. Lachner and
T. H. Eraser, National Museum of Natural
History; M.-L. Penrith, South African Mu-
seum; George R. Vliller, Tropical Atlantic
Biological Laboratory; R. S. Gaille, Texas
Parks and Wildlife Department; M. Leible,
Universidad Catolica de Chile; C. R.
Robins, University of Miami; J. Nielsen,
Universitetets Zoologiske Museum; B.
Nafpaktitis, University of Southern Cali-
fornia; T. Abe, University of Tokyo; R.
Backus and J. Craddock, Woods Hole
Oceanographic Institution; and C. Karrer,
Zoologisches Museum, Berlin. W. Klause-
witz of the Natur-Museimi Senckenberg
provided invaluable information on the
holotype of Microichthys coccoi, and E.
Tortonese of the Museo Civico di Storia
Naturale, Genoa, answered numerous ques-
tions about problematical forms such as
Pomatomichthys constanciae and Micro-
icJithys sanzoi.
I am greatly indebted to Ernst Mayr,
Giles W. Mead and Karel F. Liem for
their guidance, criticism, and support of
my work. I am also grateful to Richard L.
Haedrich for reading the manuscript and
assisting in the planning of this research.
Special thanks are extended to G. S. Myers
for assistance with taxonomic problems.
I wish to thank the staffs of the Fish
Department, Museum of Comparative
Zoology, and Department of Natinal Sci-
200 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3
ences, Boston University, for their useful
comments and practical help. Karen Green-
leaf and Pat Allen typed the final draft of
this manuscript.
Illustrations of eleven of the twelve
species of Epigonus were prepared by L.
Laszlo Meszoly. Jordan and Jordan's il-
lustration of E. fragilis (Fig. 10) was made
available through the courtesy of the
Carnegie Museum.
Finally, a hearty vielen Dank to my wife
for her patience, encouragement, and ed-
itorial assistance.
Support for this work was provided by
NSF Graduate Fellowships during 1966
to 1971 and a grant from Harvard Uni-
versity's Committee on Evolutionary Bi-
ology (GB7346).
LITERATURE CITED
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Wellington Evening Post.
Bailey, N. T. 1959. Statistical Methods in
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Barnard, K. H. 1927. A monograph of the
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Bauchot, M. L., and M. Blanc. 1961. Poissons
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Bertolini, F. 1933. Apogonidae. Fauna e flora
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BiNi, G. 1968. Atlante dei Pesci delle Coste
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Editrice. 163 pp.
Bleeker, p. 1876. Systema Percarum revisum.
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Braxjer, a. 1906. Die Tiefsee-Fische. I. Syste-
matischer Teil. Wiss. Ergeb. Deut. Tiefsee-
Exped. "Valdivia," 1898-1899, 15: 1-420.
Breder, C. M., and D. E. Rosen. 1966. Modes
of Reproduction in Fishes. Garden City,
N. Y.: Natural History Press. 941 pp.
Brown, M. E. 1957. Experimental studies on
growtli. In The Physiology of Fishes, Vol.
I, M. E. Brown (ed. ). New York: Academic
Press, Inc., pp. 361-400.
Buen, F. de. 1959. Notas preliminares sobre
la fauna marina preabismal de Chile, con
descripcion de una familia de rayas, dos
generos y siete especies nuevos. Bol. Mus.
Nac. Hist. Natur., 27(3): 171-201.
Canestrini, J. 1860. Zur Systematik der Per-
coiden. Verb. Zool. Bot. Ver. Wien, 10:
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APPENDIX
The following chart lists all meristic and
niorphometric data for the holotypcs of
E. oUiiolepis sp. nov. and E. pectinifer sp.
nov. Measurements are given in milli-
meters.
E.
oligolepis
E. pectinifer
USNM 2 077 18
USNM 207725
MERISTIC DATA
Dorsal fin
VII-I, 10
VII -I, 9
Anal fin
II, 9
II, 9
Pectoral fin
18
15
Pelvic fin
I, 5
I, 5
Lateral line
sc ale s
34
47
Gill r ake r s
31
27
Pyloric caeca
10
6
Ve r tebr ae
10 + 15
10 + 15
Pleural ribs
7
8
E p i p 1 e u r a 1 rib
s
7
6
MORPHOMETRIC
DATA
SL
90. 8
114. 3
HL
33. 2
40. 5
Body depth
21.2
28. 1
Head height
17. 1
18. 8
Eye diameter
14. 5
16. 1
Snout 1 e n g t li
7. 8
10. 9
I n t e r 0 r b i t a 1 w i
idth
8. 4
10. 0
Maxillary leng
th
15. 6
18. 2
Lower jaw len
gth
16. 3
18. 5
Caudal p c <l u n c :
d e p t h
le
8. 9
11. 7
Caudal p e d u n c :
length
le
23. 6
32. 1
D2l
10. 9
13. 7
All
11.1
13. 7
P2I
12. 3
14. 4
us ISSN 0027-4100
BulLetln OF THE
Museum of
Comparative
Zoology
The Spider Family Anyphaenidae
in America North of Mexico
NORMAN PLATNICK
HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSETTS, U.S.A.
VOLUME 146, NUMBER 4
19 SEPTEMBER 1974
PUBLICATIONS ISSUED
OR DISTRIBUTED BY THE
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HARVARD UNIVERSITY
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BuLLETiN 186a-
Memoirs 1864-1938
JoHNSONiA, Department of Mollusks, 1941-
OccAsiONAL Papers on Mollusks, 1945-
SPECIAL PUBLICATIONS.
1. Whittington, H. B., and E. D. I. Rolfe (eds.), 1963. Phylogeny and
Evolution of Crustacea. 192 pp.
2. Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredini-
dae (Mollusca: Bivalvia). 265 pp.
3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echinoderms.
284 pp.
4. Eaton, R. J. E., 1974. A Flora of Concord. 211 pp.
Other Publications.
Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine.
Reprint.
Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of
Insects.
Creighton, W. S., 1950. The Ants of North America. Reprint.
Lyman, C. P., and A. R. Dawe (eds.), 1960. Symposium on Natural
Mammalian Hibernation.
Peters' Check-list of Birds of the World, vols. 2-7, 9, 10, 12-15.
Proceedings of the Nevi' England Zoological Club 1899-1948. (Complete
sets only.)
Publications of the Boston Society of Natural History.
Price list and catalog of MCZ publications may be obtained from Publications
Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massa-
chusetts, 02138, U.S.A.
© The President and Fellows of Harvard College 1974.
THE SPIDER FAMILY ANYPHAENIDAE IN AMERICA
NORTH OF MEXICO'
NORMAN PLATNICK-
Abstract. E\'idence from the tracheal system,
claw tufts and courtship behavior is used to
justify the family status of Anyphaenidae. Sug-
gested relationships between Anyphaenidae and
Clubionidae, Amaurobiidae and Argyronetidae
are disclaimed. The faniih' Amaurobioididae is
newly synonymized with Anyphaenidae. Generic
problems within the family are discussed. The
thirty-six species occurring north of Mexico are
described, their diagnostic characters pointed out
and illustrated, their distributions mapped, notes
on their habits given, and keys to genera,
species groups and species provided. The genera
AuiipliaencUa and Ciagus are newly synonymized
witli Wulfila. Thirteen species are described as
new: Amjpliacna aJachua, A. arhida, A. uutumna,
A. catalina, A. cochise, A. gertschi, A. fiiJjIwides,
A. hespar, A. lacka, A. rita, Aijslia aninda,
Wulfila hnjantac and W. wiinda. Nineteen new
synonymies are recognized.
INTRODUCTION
This study had three objectives: to de-
termine whether or not the anyphaenids
should be treated as a distinct family; to
examine the relationships between the any-
phaenids and the other groups of spiders
with which they have been associated in
the past; and to reclassify the species oc-
' This study was presented to the Department
of Biology at Harvard University in partial ful-
fillment of the requirements for the degree of
Doctor of Pliilosophy.
"Present address: Department of Entomology,
The American Nhiseum of Natural History, Cen-
tral Park West at 79th Street, New York, New
York 10024.
curring north of Mexico on generic and
specific levels.
The anyphaenids are a diverse group
with perhaps five hundred species. Thirty-
six species are known to occur in America
north of Mexico and are included here.
About 375 species have been described
from the Neotropic region, as well as
around ten from the Palearctic and five
from the Oriental. The South American
species show the widest spectrum of body
forms; they range from 2-25 mm in length
and are often intricately colored or have
peculiarly elongate chelicerae or legs.
As in most spiders, little is known of the
ecology or behavior of anyphaenids. They
are wandering hunters. In the eastern
United States, where long-legged species
predominate, they are most often collected
by sweeping foliage in fields and meadows,
and seem to be primarily noctiu-nal. How-
ever, in the western United States, where
most species have shorter legs, they are usu-
ally found in forests by sifting through
litter and turning logs and stones. They
feed on various groups of insects, and
though they have been observed to prey
heavily on such Lepidoptera as the fall
webworm, IlijpJiantria cunea (Warren et
al., 1967), they are proliably not very se-
lective. In captivity they will consume
Drosophila eagerly. Their principal ene-
mies in nature are the mud-dauber wasps
of the family Sphecidae, as evidenced from
the hundreds of individuals, particularly
Bull. Nhis. Comp. Zool, 146(4) : 205-266, September, 1974 205
206 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
of the diurnally active genus Aijsha, that
are frequently collected from wasp nests.
Krombein ( 1967 ) cites especially the wasp
genus Trypargilum in this respect. Like
most nearctic spiders, males and females
usually mature in early spring, with males
living through early summer and females
living through the summer. In some south-
ern species, however, both sexes are found
matm-e year-round. Also, some species in
the Amjphaena celer group are matiu-e
throughout the winter. Anyphaenids make
little use of silk, other than in building re-
treats under leaves or stones and of course
in building egg sacs, which are usually
round, made of soft white silk, not leathery
or papery, and contain between 50 and
150 eggs.
The North American species are 2-9 mm
long; the largest species belong to the
genus Aijsha, the smallest to Wulfih.
There are always eight eyes in two rows;
the median eyes are usually closer to the
laterals than to each other; unlike many
gnaphosids, the eyes are always round, and
unlike many clubionids, the anterior me-
dian eyes are usually smaller than the
others. Other than the genitalia, the main
structural differences between males and
females are the sternal and coxal modifi-
cations (pointed spurs, rounded knobs, or
clumps of short thick setae) found on
males in some groups.
In many species groups it would be im-
possible to distinguish the species without
using genitalic characters. The palpus
(Text-fig. 3) usually has a large median
apophysis (the shape of which is often
species-specific), a small conductor and a
conspicuous embolus. Besides the retrolat-
eral tibial apophysis (almost always of
great diagnostic value) a ventral tibial
apophysis (some Aijsha) or a retrolateral
patellar apophysis (some Teudis) may be
present. The female epigyna and internal
genitalia are extremely diverse and diffi-
cult to characterize. The two epigynal
openings are located posteriorly and are
extremely difficult to see unless a portion
of the male embolus has been left behind
after mating. Many species have an addi-
tional anterior median epigynal opening
into which the retrolateral tibial apophysis
or median apophysis fits during mating.
The genitalia of anyphaenids, particularly
of the South American species, are more
complex than those of clubionids and
gnaphosids. Among the clubionids, only
Chiracanthium has genitalia that seem in
any way close to those of anyphaenids.
For the area treated here, only three im-
portant papers have been published on
anyphaenids. Bryant (1931) summarized
the very sparse data then available on the
group in the United States, while Chicker-
ing (1937, 1940) described many of the
species occurring in Panama and the Canal
Zone, a number of which also occur in the
United States.
ACKNOWLEDGEMENTS
I would like to thank first Herbert W.
Levi for his painstaking and patient help
with all aspects of this project. Willis
Gertsch contributed much of his knowl-
edge of the group as well as the drawings,
done by the late Wilson Ivie, of the genus
Oxijsorna.
This investigation was supported in part
by Public Health Service Research Grant
AI-01944 from the National Institutes of
Allergy and Infectious Diseases, H. W.
Levi, principal investigator; by Grant GB-
36161 from the National Science Founda-
tion, H. W\ Levi, principal investigator;
and by Grant GB-19922 from the National
Science Foundation, R. C. Rollins, princi-
pal investigator. The Department of Biol-
ogy, Harvard University, by means of
Summer Research Grants in Evolutionary
Biology in 1971 and 1972 and a Richmond
Fellowship in 1973, afforded me much of
the time and field work necessary to com-
plete this work. Miss Suzanne Barbier of
Radcliffe Gollege assisted greatly with the
examination of tracheal systems and her
work is deeply appreciated.
Finally, the following people loaned
SpTDKn FA\rii,v ANVPiiAEXinAK • Platnirk
207
specimens from their private collections
or from their cited institntions: Paul II.
Aniaud, Jr. (California Academy of Sci-
ences), Joseph A. Beatty, Jr., James E.
Carico, John A. L. Cooke (American Mu-
seum of Natural History), Charles D.
Dondale (Canadian National Collections),
R. R. Forster (Otago Museum), Willis J.
Gertsch (American Museum of Natural
History), Al Jung, B. J. Kaston, Robin
Leech, William B. Peck, Vince Roth, Rich-
ard J. Sauer (Michigan State University),
William A. Shear, Bea Vogel, H. K. Wal-
lace, Fred R. Wanless (British Museum,
Natiu-al History), and Howard V. Weems
(Florida State Collection of Arthropods).
THE FAMILY STATUS OF
ANYPHAENIDAE
Simon considered tlie anyphaenids to be
a subfamily of the large family Clubionidae
and used as the key character for distin-
guishing the anyphaenids the advanced
placement of the tracheal spiracle. Later
authors, notably Petrunkevitch and Bris-
towe, thought this character so significant
that they gave the anyphaenids family sta-
tus, though still believing the group to be
closely related to the Clubionidae. The
comparatively recent discovery that in
some families closely related, congeneric
species sometimes have very different res-
piratory systems (see Levi, 1967) has led
most arachnologists to denigrate the im-
portance of respiratory structm-es as macro-
taxonomic characters. Thus most modern
arachnological works still treat the any-
phaenids as a subfamily of Clubionidae.
A notable exception, however, is Lehtinen
( 1967 ) , who maintains ( correctly, I be-
lieve) that the classical family Clubionidae
is a highly polyphyletic assemblage of un-
related two-clawed spiders that lack any
noticeable modifications of the body. Leh-
tinen splits the clubionids into several fam-
ilies, largely but not strictly along the lines
of the old subfamily divisions, and accords
the anyphaenids full status as a family.
Forster (1970) agrees with this assessment
of the anyphaenids.
To check on the validity of this classifi-
cation, a variety of clubionid genera were
examined and compared with anyphaenids,
with the result that the anyphaenids arc
here considered a distinct family, for two
major reasons. One is the classical reason
— the tracheal system. Examination of the
tracheae of males and females of the club-
ionids CAuhiona o])esa Hentz, Chiracan-
thium mildei L. Koch, Trachelus tratujuil-
his- (Hentz), Castkineira cin<i,uluta (C. L.
Koch), Agroeca pratensis Emerton, Phniro-
timpus alarius (Hentz), and the any-
phaenids Am/pJmena celer (Hentz), Amj-
phaena pectorosa L. Koch, Amjphaena
calif ornica ( Banks ) and Aijsha <i,racilis
(Hentz) disclosed three major differences
between anyphaenid and clubionid tra-
cheae (see Methods for the technique
used). First, anyphaenid tracheae extend
through the pedicel of the spider into the
cephalothorax and legs, while those of
clubionids are restricted to the abdomen
(see Figs. 47 and 50). Associated with this
is the externally observable advanced
placement of the tracheal spiracle in any-
phaenids. Second, the tiacheae are rela-
tively much larger in anyphaenids. In all
the clubionids examined, even the main
tracheal tubes are very thin and narrow;
anyphaenid tracheae are three to four
times as wide. Third, none of the clubionid
species examined showed any sexual di-
morphism in the tracheal system, whereas
male anyphaenids have considerably larger
tracheae than do the females. The size
of the tracheae may be correlated with the
high activity levels of anyphaenids: my
collecting experience indicates that they
can run extremely rapidly when disturbed.
The larger size of the tracheae in males
may be associated with the increased respi-
ration necessary for the extra activity re-
(juired to locate, court and copulate with
a female. Anyphaenid courtship is (.ex-
tremely active; films of the courtship of
Anijphacna accentiiata show that the abdo-
208 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
embolus
median apophysis
conductor
retroloteral tibial
apophysis (RTA)
Text-Figures 1-3. Claw tuft of Clubiona obesa Hentz, lateral view, diagrammatic. 2. Claw tuft of Aystia gracilis
(Hentz), lateral view, diagrammatic. 3. Generalized palpal structure of Anyphaena.
men of the male is vibrated up and down
so rapidly that only a blur is visible
(Thompson, G. H., and E. R. Skinner,
Courtship in Spiders, Oxford Scientific
Films). Although the mating behavior of
very few species in either group has been
studied in detail, the vast difference be-
tween anyphaenid courtship and the rather
sluggish courtship behavior of Clubiona
and related genera wovild seem to provide
additional evidence for separating the two
groups (Platnick, 1971).
Evidence that is probably just as impor-
tant as the tracheae for considering Any-
phaenidae a distinct group is provided by
the claw tufts. Clubionids have claw tufts
that are composed of numerous straight
simple setae densely clumped together
( Text-fig. 1 ) . Anyphaenid claw tufts, how-
ever, are composed of two rows of large,
lamelliform setae that are greatly expanded
at their distal ends (Text-fig. 2). All the
anyphaenids examined have these peculiar,
easily recognizable claw tufts, but so far
as known, no clubionids do, though some
phruroliths have superficially similar claw
tufts.
For these reasons, Anyphaenidae is here
considered a distinct family not very
closely related to any of the groups cur-
rently included in the Clubionidae.
RELATIONSHIPS OF THE FAMILY
ANYPHAENIDAE
In addition to the clubionids, the any-
phaenids have been associated with three
other families of spiders: Amaurobiidae,
Argyronetidae and Amaurobioididae. Leh-
tinen (1967) placed the anyphaenids in
his branch Amaurobiides and stated that
they are probably derived from Amauro-
biidae: Macrobuninae and therefore lack
ecribellate, two-clawed relatives. Forster
( 1970 ) agreed with the placement of Any-
phaenidae in Amaurobiides (and specifi-
Spider Family Axypiiaenidae • Plalnick
209
cally included the family in his supeifainily
Dictynoidca) but cited the families Ar-
gyionetidae and Amaurobioididae as close
relatives. Representatives of all three fami-
lies were examined to determine the degree
of their relationship, if any, to the Any-
phaenidae.
Lehtinen gave no evidence for his state-
ment that the anyphaenids are probably
deri\'atives of Amaurobiidae: Macrobuni-
nae, presumably because there seems to be
none. An examination of specimens of
one genus in this subfamily, Arctohius
Lehtinen, indicates that it would be diffi-
cult to find araneomorph spiders less likely
to have given rise to the anyphaenids. The
species of Arctohius are cribellate, three-
clawed spiders that lack claw tufts and
possess an unelaborated tracheal system.
Further, the genitalia show no similarities
to those of anyphaenids.
Likewise, Forster gave no evidence for
associating the family Argyronetidae with
tlie an\q3haenids; his decision to do so was
based, I believe, on the similarities in the
tracheal systems of the two groups. The
elaboration of the tracheal system in Ar-
gyroneta, however, is probably associated
with their invasion of an aquatic habitat
and the resultant demands on the respira-
tory system. All the other characters, in-
cluding the three claws, lack of claw tufts
and the characteristic pattern of ti'icho-
bothria distribution, indicate that Argt/ro-
neta is, as it is usually regarded, a close
relative (if not actually a member) of the
family Agelenidae.
The family Amaurobioididae was cre-
ated by Hickman ( 1949 ) for the single
genus Ammirohioides O. P. -Cambridge,
which has at various times been included
in the families Drassidae ( = Gnaphosi-
dae), Ctenidac, Clubionidae and Miturgi-
dae. The genus is known from New Zea-
land, Tasmania, southern Chile and South
Africa. The spiders live in rock crevices
in the tidal zone, where they build tubular
silk retrtnits and are regularly submerged
at high tide (Lamoral, 1968).
Specimens of this rare genus provided by
R. R. Forster revealed not only a typically
anyphac>nid-]ike tracheal system, but also
the lamelliform claw tufts so characteristic
of anyphaenids. Further, the genitalia are
close to those of the anyphaenid genus
Oxysouui, and the body form is similar to
that of several species of anyphaenids
known from Chile, Peru, and Argentina.
For these reasons, the family Amauro-
bioididae is newly synonymized with
Anyphaenidae in the taxonomic section of
this paper.
Thus the problem of the correct macro-
taxonomic placement of Anyphaenidae has
been clarified but not solved by this study
of the groups with which the family has
been associated in the past. Futiu-o work
should start with an examination of the
family Miturgidae (as construed by Leht-
inen ) .
Although it was necessary to limit the
scope of the detailed revision to the man-
ageable number of species occurring north
of Mexico, all available specimens from
other areas were examined to gain an
overview of the family. Preliminary im-
pressions indicate that the family probably
originated in the southern half of South
America with subsequent radiations north-
ward. As indicated by the ability of
Amaurohioides to withstand prolonged sub-
mersion, it is likely that early anyphaenids
were able to survive hydrochore dispersal
by rafting, etc., across considerable ex-
panses of water.
GENERIC PROBLEMS IN THE
ANYPHAENIDAE
The generic taxonomy of anyphaenids
is currently chaotic. Every author who has
worked with the group, including Petrunke-
vitch (1930), Bryant (1931) and Chicker-
ing (1937), has expres.sed frustration at
the confusion and ambiguity in the use of
many of the most common generic names.
One of the principal causes of this con-
fusion is the interesting e\olutionary pat-
tern encountered time and again witliin
210 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
this family: species tend to occur in groups
that are remarkably homogeneovis in
genitalic structure but quite distinct from
other such groups. Often many of the
species in these groups are sympatric, are
found in a rather limited area and are
clearly the result of radiation within that
area. An excellent example of this is the
occurrence of nine closely related species
of the Anyphaena celer species group in
the mountains of southeastern Arizona. It
is tempting to consider each of these
groups a genus, as unambiguous key char-
acters are then available to distinguish
genera. Such an approach would at least
double the number of genera found in the
United States, and, if applied to the Central
and South American fauna, would neces-
sitate the creation of a vast number of
new genera. If, instead, characters refer-
ring to the general body form are used, a
more workable classification in terms of
both number and size of genera results.
Unfortunately, this makes the unambiguous
definition of genera much more difficult
and makes keys to genera awkward and
cumbersome. With either approach, how-
ever, reliable genera composed of mo-
nophyletic groups of species can be estab-
lished.
The second approach to anyphaenid clas-
sification has been taken by the majority
of former authors, and is continued in this
work. Thus the European genus Any-
phaena is used for the bulk of the any-
phaenids occurring in the United States,
even though only one of our species, Any-
phaena aperta, is actually a close relative
of the European Anyphaena accentuata,
type species of the genvis. Nonetheless,
all the species here included in Anyphaena
share a basic body form. The neotropical
genus Wulfila is used for all the pale,
long-legged species, even though they are
genitalically quite diverse; the other genera
used here are similarly construed. Although
this system is not wholly satisfactory, it
seems decidedly better than creating a
host of new generic names that are likely
to fall into synonymy when a detailed ge-
neric revision of the group as a whole can
be carried out.
METHODS
Tracheae were examined by dissecting
away the dorsal cuticle of the abdomen
and boiling the spider in ten percent
sodium hydroxide for ten minutes. By this
method, all the soft structures in the
abdomen are digested away, leaving the
tracheae intact.
Types of the new species are being
deposited in the American Museum of
Natural History, New York City, and the
Museum of Comparative Zoology, Harvard
University. Type depositories are abbrevi-
ated as follows: AMNH — American Mu-
seum of Natural History, BMNH — British
Museum, Natural History, MCZ — Museum
of Comparative Zoology.
Measurements and drawings were made
with a standard ocular grid. Measurements
of gross morphological featiu-es are ac-
curate to ~ 0.04 mm; measurements of
ocular featiu'es are accurate to ~ 0.01 mm.
Rather than selecting a small number of
measurements and providing means and
standard deviations for these on the basis
of a small series of specimens, one male
and one female of each species were
measured in detail. As only one of the
species included here shows any significant
variation in size, this procedure was deemed
more informative. Actual measurements
are given rather than ratios since in many
cases (e.g., Anyphaena catalina and A.
arhida) closely related species differ sig-
nificantly in size but not in their relative
proportions. Most of the measurements
taken are self-explanatory, though a few
need furdier comment. Cephahc width
refers to the width of the carapace at a
point just behind the posterior median
eyes, and thus provides an indication of
the degree to which the carapace is nar-
rowed in front.
The difficult problem of accurately de-
scribing die eye relationships has been
Spider Family Anyphaenidae • Platnick 211
solved by providing a set of measurements
from wliich it is possible to reconstrnct,
using grapli paper, the exact eye arrange-
ment. Diameters are given using the con-
ventional abbreviations (AME = anterior
median eye, ALE = anterior lateral eye,
PME = posterior median eye, PLE =
posterior lateral eye). The length of each
eye row is measured from the lateral edge
of one lateral eye to the lateral edge of
tlie other lateral eye. Curvature of the eye
rows is described as viewed frontally, not
dorsally. This was accomplished by posi-
tioning the spider in sand, a technique
found most useful for making all the
measurements. The dimensions of the
median ocular quadrangle (MOQ) are
given, as well as the distances between
each of the eyes. The latter measurements
extend between the edges of the lenses of
the eyes under consideration (not just
between the dark circles surrounding each
eye).
The relative length and thickness of each
leg is indicated by the tibial length index —
the tibial width divided by the tibial
length, with the result multiplied by 100
to obtain a whole number. All tibial
measurements were taken from a dorsal
view and refer to the maximum lengths
and widths. The lower the tibial index,
the longer and thinner the leg; conversely,
the higher the index, the shorter and thicker
the leg. In practice the index varies from
around 3 to 35.
Ventral spination of the leg segments is
indicated by the standard formula in which
the number of spines on the proximal,
median and distal thirds of the leg segment
are given. Only ventral spines, not lateral
ones, are included, and any even number
in the formula may be taken to represent
a pair of spines. Unless the last number is
followed by an asterisk, the last pair of
spines is terminally located. Thus, for
example, the formula 2-2-2* indicates that
the segment bears three pairs of ventral
spines, the last pair of which is not termi-
nally located. The term "spine" is used in
its conventional arachnological sense and
rc'fers to the moxable macrosetae found
on the legs. Similarly, the term "clypeus"
is used to refer to the area between the
anterior eye row and the anterior edge of
tlie carapace and not to the small sclerite
folded under the carapace. Since neither
usage of tht> term reflects certain knowledge
of homology with the insect clypeus, the
old and established usage should be main-
tained.
Scale lines for the drawings always equal
0.1 mm. Each scale line applies to all
consecutively numbered drawings imtil a
new scale line appears. Exceptions are
noted in the captions.
TAXONOMY
Anyphaenidae
Anyphaenidae Bertkau, 1878, Arch. Naturg., 44:
358, 379. Tvpe genus Anyphaena Sundevall,
1833.
Amaurobioididae Hickman, 1949, Pap. Proc. Roy.
Soc. Tasmania, 1948: 31. Tvpe genus A77iauro-
hioides O.P.-Cambridge, 1883. NEW SYN-
ONYMY.
Diagnosis. The combination of the ad-
vanced tracheal spiracle and the lamelli-
form claw tufts will serve to distinguish the
anyphaenids from all other families.
Description. Chelicerae diaxial, not fused
together at base. Labium free. Without
cribellum or calamistrum. With one pair
of book lungs and a tracheal spiracle lo-
cated considerably anterior to the spin-
nerets, most often midway between spin-
nerets and epigastric furrow, sometimes
closer to one or the other. Eight eyes in
two rows. Six spinnerets, anterior spin-
nerets approximate, colulus represented
only by hairs, anal tubercle unmodified.
Legs prograde, metatarsi and tarsi I and
II scopulate, tarsi with two toothed claws
and claw tufts composed of lamelliform
setae.
Key to Genera
IN America north of Mexico
la. Tracheal spiracle luucli closer to epigastric
furrow tlian to spinnerets ..Aysha
212 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
lb. Tracheal spiracle roughly midway between
epigastric furrow and spinnerets 2
2a. Legs very long and thin. Leg I greatly
elongated, tibial index (width/length X
100) usually 5 or less Wulfila
2b. Legs normal, tibial index of leg I usually
8 or more 3
3a. Chelicerae with 2 retromarginal teeth
Oxysoma
3b. Chelicerae with 4-9 retromarginal denticles
4
4a. Carapace usually with two dark paramedian
longitudinal bands; chelicerae not produced
forward; femora not much darker than
other leg segments Anijphaena
4b. Carapace without dark paramedian longi-
tudinal bands; either chelicerae produced
forward or femora much darker than other
leg segments Teudis
Anyphaena Sundevall
Amjphaena Sundevall, 1833, Conspectus Arachn.,
28. Type species by monotypy Aranea ac-
centuata Walckenaer, 1802.
Diagnosis. The combination of the fol-
lowing characters will serve to distinguish
the genus in America north of Mexico:
trachael spiracle roughly midway between
epigastric furrow and spinnerets, leg I not
greatly elongated, chelicerae with 4-9
retromarginal denticles and not produced
forward, femora not much darker than
other leg segments. The carapace usually
has two dark paramedian longitudinal
bands. The genus is used here in a very
broad sense; this prevents simple diagnosis,
and makes detailed descriptions of each
species group more meaningful than a
description of the whole genus.
Uncertain names. Types of the follow-
ing species were unavailable and are too
poorly described to permit identification:
Cluhiona agresiis Hentz, 1847, type de-
stroyed; Chihiona fallens Hentz, 1847, type
destroyed, Cluhiomi suhlurida Hentz, 1847,
type destroyed; Amjphaena argentata
Becker, 1879, type lost; and Amjphaena
striata Becker, 1879, type lost. The three
Hentz Cluhiona species were transferred to
Anijphaeiui by Marx (1890), but there is
little justification for this in the vague
descriptions. All the above names are
regarded as nomina cluhia.
Species groups. Although there seem to
be several species groups of Amjphaena in
the Neotropic region, only four occur north
of Mexico. The celer group is the largest;
it has representatives at least as far south
as Panama and probably contains over
thirty species. The pectorosa and pacifica
groups are closely related and occur com-
monly in Mexico as well as the United
States; it is difficult to place females in
one group or the other unless the male is
also known; they probably contain together
at least twenty species. The accentuata
group is predominantly Palearctic and prob-
ably contains at least five species.
Key to Species Groups
la. Metatarsi I and II with one pair of ventral
spines accentuata group
lb. Metatarsi I and II with two pairs of ventral
spines 2
2a. Retrolateral tibial apophysis of males bifid,
with ventral prong elongated ( Figs. 18-20,
25-32). Epigynum of females with a hood
(Figs. 21, 23, 33, 36, 37, 39-42) ._ celer group
2b. Retrolateral tibial apophysis of males not
bifid or elongated (Figs. 55-58, 69-71).
Epigynum without a hood (Figs. 66, 67,
72, 74, 77, 79) 3
3a. Eastern United States. Coxae III and IV
of males with pointed spins (Figs. 59-62).
Female epigyna on broad sclerotized plates
(Figs. 74, 77, 79); internal genitalia lacking
long ducts (Figs. 75, 78, 80)
pectorosa group
3b. Western United States. Coxae III and IV
of males without pointed spurs, though
rounded knobs may be present. Female
epig>'na not on broad sclerotized plates
(Figs. 66, 67, 72); internal genitalia with
long, sometimes coiling, ducts (Figs. 68,
73, 76) _._. pacifica group
Anyphaena celer Group
Diagnosis. Males of the celer group may
be recognized by their retrolateral tibial
apophysis, which is usually bifid with an
elongated ventral prong (Figs. 18, 20, 26).
Females have a characteristic epigynum
consisting of a hood, two sidepieces and a
midpiece (Figs. 9, 33), though the mid-
SpiDEii Family Axyphakxidae • Pkilnick 213
piece is reduced in A. crebrispimi ;ind A.
(lixiana (Figs. 21, 23).
Description. Total length 3-7 nnn, with
males of most species between 3.3-4.6 mm,
females of most species between 4.1-5.9
mm. Carapace longer than wide, narrowed
in front to less than half its maximum
w idth. Clypens height greater than anterior
median eye diameter. Posterior median,
posterior lateral and anterior lateral eyes
subeqiial in size, larger than anterior
medians. Procurved posterior eye row
longer than recin'\'ed anterior row. Median
ocular ({nadrangle longer than wide in
front, wider than long in back. Anterior
median eyes separated by their diameter,
by their radius from anterior laterals.
Posterior medians separated by their diam-
eter, slightly closer to posterior laterals
than to each other. Anterior laterals
separated by their radius from posterior
laterals. Sternum longer than wide, un-
modified. Chelicerae with 4-5 promarginal
teeth and 6-9 reti-omarginal denticles.
Abdomen longer than wide, tracheal
spiracle midway between epigastric fur-
row and base of spinnerets. Leg formula
1423. Metatarsi I and II with two pairs
of \'entral spines. Males often with femur
III thickened distally, set with stiff short
setae ventrally; tibia III ventral spines
thickened, cone-like; coxae set with clumps
of stiff short setae. Palpus with an elon-
gated median apophysis, retrolateral teg-
ular apophysis, conspicuous curving em-
bolus and conductor. Retrolateral tibial
apophysis bifid, with dorsal prong reduced
in some species. Epigynum with hood,
two sidepieces and midpiece; two simple
spermathecae.
Variation. None of the species in this
group show any significant individual or
geographic intraspecific variation in struc-
tin-e, size or coloration.
Key to Species
la. Dorsal and \entral prongs of retrolateral
tibial apophysis ( RTA ) roughly equal in
length (Figs. 18, 19); epigynal hood wide,
more than four times the minimum width
of epigynal sidepiece ( Figs. 9, 11); east-
ern U.S 2
II). Ventral prong of retrolateral tibial apophy-
sis ( RTA ) nuieh longer than dorsal prong
(as in Figs. 26, 27); epigynal hood
narrow, less than four times the minimum
width of epigynal sidepiece ( as in Figs.
33, 36); western U.S. 3
2a. Dorsal prong of RTA broad, with a trans-
lucent ridge (Fig. 18); epigynal hood a
thick oval, sidepieces straight (Fig. 9)
— - - — _.. celer
21). Dorsal prong of RTA narrow, without a
translucent ridge (Fig. 19); epigynal hood
a thin oval, sidepieces rounded ( Fig. 1 1 )
- - _ inaculata
3a. Base of RTA expanded into a broad
triangle (Fig. 20); retrolateral tegular
apophysis prolonged medially (Fig. 3);
epigynal sidepieces more than three times
the width of epigynal hood (Fig. 21)
- crebrispina
3b. Base of RTA not expanded; retrolateral
tegular apophysis not prolonged medialK ;
epigynal sidepieces less than three times
the width of epigynal hood 4
4a. Dorsal prong of RTA bearing a sharp spur
(Fig. 25); epigynal midpiece greatly re-
duced, sidepieces widely separated pos-
teriorly (Fig. 23) dixiana
4b. Dorsal prong of RTA without a spur;
epigynal midpiece conspicuous, sidepieces
approximate posteriorly 5
5a. Males '. 6
5b. Females 14
6a. Dorsal prong of RTA with two triangular
processes separated by a concave notch
( Fig. 26 ) judicata
6b. Dorsal prong of RTA witliout triangular
processes „. 7
7a. Dorsal prong of RTA with a long recurved
hook (Fig. 29) autumna
7b. Dorsal prong of RTA without a long re-
curved hook 8
8a. Dorsal prong of RTA witli a basal hook
(Figs. 31, 38) _.-.. 9
8b. Dorsal prong of RTA without a basal
hook 10
9a. Conductor and retrolateral tegular apophy-
sis recurved (Fig. 15) catalina
9b. Conductor and retrolateral tegular apoph-
ysis not recurved ( Fig. 17 ) arhida
10a. Dorsal prong of RTA a shaiply pointed
spike ( Fig. 32 ) liespar
101). Dorsal prong of RTA not a sharply
pointed spike 1 1
11a. Fmbolus with a conspicuous enlargement
(Figs. 7, 13) .12
111). Embolus without a con.spicuous enlarge-
ment (Figs. 21, 33) 13
214 Bulletin Museum of Comparative Zoolog,ij, Vol. 146, No. 4
J?
I ^J'', ,^-^ m-p.
Anyphaeno maculota \ q\ —>>
o-
I -*
J
\ \A-
1
1
^~« ',
>-:-:
Anyphoena crebrispino
/—
\\v
Anyphaena dixiana j»- \
v^\ \
Anyphaena rita '
V f
1 ~T
1 1
\h~^
' 1
Anyphaena
cochise V
Map 1. Distributions of Anyphaena arbida, A. autumna, A. catalina, A. celer, A. cochise, A. crebrispina, A. dixi-
ana, A. gibboides, A. hespar, A. judicata, A. maculata, A. marginalis and A. rita.
12a. Dorsal prong of RTA more than half the
length of ventral prong ( Fig. 35 )
cochise
12b. Dorsal prong of RTA less than half the
length of ventral prong (Fig. 28)
rita
13a. Median apophysis sharply pointed; con-
ductor short, bent (Fig. 14); Oregon and
Utah gibboides
13b. Median apophysis rounded; conductor
long, straight (Fig. 6); Arizona and New
Mexico marginalis
14a. Epigynal hood wider than long; midpiece
not wider than hood, without constric-
tions; sidepieces very wide (Fig. 40);
Oregon and Utah gibboides
14b. Epigynal hood as long as wide or midpiece
wider than hood or sidepieces narrow;
Arizona and New Mexico 15
15a. Epigynal midpiece a very broad triangle
( Fig. 37 ) rita
15b. Epigynal midpiece othenvise 16
16a. Spermathecae much further apart pos-
teriorly than anteriorly ( Fig. 49 ) _-, hespar
16b. Spemiathecae as far apart anteriorly as
posteriorly 17
17a. Epigynal hood much wider than long ( Fig.
39 ) _ autumna
17b. Epigynal hood as long as wide —18
18a. Epigynal midpiece less than twice the
length of epigynal hood (Fig. 41)
catalina
18b. Epigynal midpiece more than twice the
length of epigynal hood ...19
19a. Epigynal midpiece a short triangle (Fig.
33 ) judicata
19b. Epigynal midpiece an elongate triangle
(Fig. 36) marginalis
Anyphaena celer (Hentz)
Map 1; Figures 1, 9, 10, 18
Chibiona celer Hentz, 1847, J. Boston Soc. Natur.
Hist., 5: 452, pi. 23, fig. 20 ( 9 ). Male holo-
type, female allotype from Alabama and North
Carolina in the Boston Soc. Natur. Hist.
(Boston Museum of Science), destroyed by
beetles.
Anyphaena incerta Keyserling, 1887, Verb. zool.
hot. Ces. Wien, .37: 452, pi. 6, fig. 22 ( $ ).
Female holotype from Cambridge, Massachu-
setts, in MCZ, examined. Emerton, 1890, Trans.
Connecticut Acad. Sci., 8: 186, pi. 6, figs.
2-2d, $,9.
Anyphaena celer, Simon, 1897, Hist. Natur.
Araign., 2: 96. Bryant, 1931, Psyche, 38: 111,
pi. 6, fig. 9, pi. 8, figs. 25, 28, $, 9. Chickering,
1939, Pap. Michigan Acad. Sci., 24: 51, figs.
Spider Family Axyphakxidae • PJaliiick 215
Plate 1
Figures 1-8. Left palpi, ventral view. Figures 9. 11. Epigyna, ventral view. Figures 10, 12. Internal genitalia,
dorsal view. 1,9,10. /Anyphaena ce/er (Hentz). 2,11,12. Anyphaena maculata (Banks). 3. Anyphaena crebri-
sp'ma Chamberlin. 4. Anyphaena dixiana (Chamberlin and Woodbury). 5. Anyphaena judicata O. P. -Cambridge.
6. Anyphaena marginalis (Banks). 7. Anyphaena rita new species. 8. Anyphaena autumna new species.
216 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
t>'
1-4, $, 9. Comstock, 1940, Spicier Book,
rev. ed., p. 577, figs. 634-6.35, $,9. Kaston,
1948, Bull. Connecticut Geol. Natur. Hist.
Surv., 70: 407, figs. 1471-1476, $,9. Roewer,
1954, Katalog der Araneae, 2:528. Bonnet,
1955, Bibliographia Araneorum, 2: 343.
Gatjenna celer, Comstock, 1912, Spider Book, p.
563, figs. 634-635, $,9.
Dia<ino.sis. Anyphaena celer is most
closely related to A. maculata. Males of
both species have dorsal and venti-al RTA
prongs roughly equal in length, but A. celer
males may be distinguished by the trans-
lucent ridge on their dorsal prong (Fig.
18). Females may be separated by the
straight epigynal sidepieces and widely
oval epigynal hood of A. celer (Fig. 9).
Male (Jackson Co., Illinois). Total
length 4.54 mm. Carapace 2.12 mm long
1.58 mm wide, cephalic width 0.83 mm
clypeus height 0.07 mm, pale yellow with
thin dark broken border and two dark
paramedian longitudinal bands. Eyes:
diameters (mm): AME 0.06, ALE 0.12,
PME 0.11, PLE 0.12; anterior eye row
0.44 mm long, slightly recurved; pos-
terior eye row 0.60 mm long, procurved;
MOQ length 0.24 mm, front width 0.19
mm, back width 0.31 mm; eye interdis-
tances (mm): AME-AME 0.06, AME-
ALE 0.03, PME-PME 0.10, PME-PLE
0.10, ALE-PLE 0.05.
Sternum 1.06 mm long, 0.94 mm wide,
pale yellow with dark markings opposite
coxae, translucent border and darkened
extensions to coxae. Chelicerae 0.79 mm
long with 4 promarginal teeth and 8 retro-
marginal denticles, pale yellow with boss
outlined in gray. Labium and endites pale
yellow, darkest proximally; endites not
invaginated.
Abdomen 2.30 mm long, 1.62 mm wide,
pale white with transverse rows of dark
markings; venter with scattered dark mark-
ings. Epigastric furrow 0.85 mm from
tracheal spiracle, spiracle 0.88 mm from
base of spinnerets.
Legs pale yellow with scattered dark
markings. Tibial lengths (mm) and
indices: I 1.98, 12; II 1.82, 14; III 1.17, 21;
IV 1.73, 16. Ventral spination: tibiae I-IV
2-2-2; metatarsi I, II 2-2-0, III 2-0-2, IV
2-2-2. Femur III thickened distally with
clump of short thick setae ventrally. Tibia
III ventral spines 1, 2 on retrolateral side
thickened, cone-like. Coxae III, IV pro-
lateral ventral surface with clump of short
thick setae.
Palpus as in Figures 1, 18.
Female (Wayne Co., Ohio). Coloration
as in male. Total length 5.87 mm. Carapace
2.07 mm long, 1.39 mm wide, cephalic
width 0.86 mm, clypeus height 0.05 mm.
Eyes: diameters (mm): AME 0.05, ALE
0.10, PME 0.10, PLE 0.10; anterior eye row
0.42 mm long, recurved; posterior eye row
0.58 mm long, procurved; MOQ length
0.30 mm, front width 0.19 mm, back width
0.32 mm; eye interdistances (mm): AME-
AxME 0.07, AME-ALE 0.05, PME-PME
0.12, PME-PLE 0.07, ALE-PLE 0.08.
Sternum 0.99 mm long, 0.88 mm wide.
Chelicerae 0.76 mm long with teeth as in
male.
Abdomen 4.10 mm long, 2.13 mm wide.
Epigastric furrow 1.57 mm from tracheal
spiracle, spiracle 1.37 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 1.58, 15; II 1.42, 16; III
1.01, 24; IV 1.60, 15. Ventral spination:
tibiae I, II 2-2-2, III, IV 1-1-2; meta-
tarsi I, II 2-2-0, III 2-0-2, IV 2-2-2.
Epigynum as in Figure 9, internal geni-
talia as in Figure 10.
Natural history. Mature males have been
taken every month except June, mature
females year-round. Specimens have been
taken in houses, deciduous forests, on
leaves, flowers, ti-eesides, in pitfalls and
footprints in snow.
Distribution. Eastern United States from
southern New England west to Wisconsin,
south to Florida and Texas (Map 1).
Anyphaena maculata (Banks)
Map 1; Figures 2, 11, 12, 19
Caijcnna maculata Banks, 1896, Trans. Amer.
Ent. Soc, 23: 64. Male holotype from Wash-
ington, D.C., in MCZ, examined. Bishop and
Spider Family Anyphaenidak • Plat nick 217
N^^
Figures 13-17. Left palpi, ventral view,
ventral view. Figures 22, 24. Interna
Anyphaena gibboides new species. 15.
17. Anyphaena arbida new species. 18
22. Anyphaena crebrispina Channberlin.
Plate 2
Figures 18-20. Left palpi, retrolateral view. Figures 21, 23. Epigyna,
genitalia, dorsal view. 13. Anyphaena cochise new species. 14.
Anyphaena catalina new species. 16. Anyphaena hespar new species.
Anyphaena celer (Hentz). 19. Anyphaena maculata (Banks). 20. 21,
23, 24. Anyphaena dixlana (Chamberlin and Woodbury).
218 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
Plate 3
Figures 25-32. Left palpi, retrolateral view. Figure 33. Epigynum, ventral view. Figure 34. Internal genitalia,
dorsal view. 25. Anyphaena dixiana (Chamberlin and Woodbury). 26, 33, 34. Anyphaena judicata O. P. -Cam-
bridge. 27. Anyphaena marginalis (Banks). 28. Anyphaena rita new species. 29. Anyphaena autumna new
species. 30. Anyphaena gibboides new species. 31. Anyphaena catalina new species. 32. Anyphaena hespar
new species.
Spider Family Anyphaenidae • Platnick 219
Crosby, 1926, T. Elislia Mitclioll Sci. Soc, 41:
189, pi. 24, figs. 37, 38, $, 9.
Amjphacna nuiciihita, Simon, 1897, Hist. Natiir.
Araign., 2: 96. Brvant, 1931, Psyche, 38: 111,
pi. 6, fig. 8, pi. 8, fig. 31, $, 9. Kaston, 1948,
Bull. Connecticut Geol. Natur. Hist. Surv.,
70: 409, figs. 1457-1458, $,9. Roewer, 1954,
Katalog der Araneae, 2: 529. Bonnet, 1955,
Bibliographia Araneoruni, 2: 345.
Diagnosis. Anyphoena inaculata is most
closely related to A. celer. Males may be
distinguished by the short dorsal prong of
the RTA, which lacks a translucent ridge
(Figure 19); females by their rounded
epigynal sidepieces and narrowly oval
epigynal hood (Figure 11).
Male (Durham Co., North Carolina).
Coloration as in Amjphaena celer. Total
length 3.74 mm. Carapace 2.09 mm long,
1.54 mm wide, cephalic width 0.77 mm,
clvpeus height 0.08 mm. Eyes: diameters
(mm): AME 0.07, ALE 0.11, PME 0.10,
PLE 0.10; anterior eye row 0.44 mm long,
recurved; posterior eye row 0.58 mm long,
procurved; MOQ length 0.23 mm, front
width 0.20 mm, back width 0.31 mm; eye
interdistances (mm): AME-AME 0.05,
AME-ALE 0.03, PME-PME 0.10, PME-
PLE 0.10, ALE-PLE 0.04.
Sternum 1.08 mm long, 0.79 mm wide.
Chelicerae 0.63 mm long with 4 promar-
ginal teeth and 8 retromarginal denticles.
Abdomen 2.02 mm long, 1.08 mm wide.
Epigastric furrow 0.31 mm from tracheal
spiracle, spiracle 0.41 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 2.00,
11; II 1.75, 13; III 1.12, 24; IV 1.82, 15.
Ventral spination: tibiae I, II 2-2-2*, III,
IV 2-2-2; metatarsi I 2-1-0, II 2-2-0, III
2-0-2, IV 2-2-2. Modifications of third
leg as in A. celer.
Palpus as in Figures 2, 19.
Female (Pope Co., Illinois). Coloration
as in male of A. celer.
Total length 4.68 mm. Carapace 2.07
mm long, 1.60 mm wide, cephalic width
0.97 mm, clypeus height 0.08 mm. Eyes:
diameters (mm): AME 0.08, ALE 0.10,
PME 0.10, PLE 0.11; anterior eye row 0.48
mm long, recurved; posterior eye row 0.63
mm long, procur\'ed; MOQ length 0.30
mm, front width 0.22 mm, back width 0.33
nun; eye interdistances (mm): AME-
AME 0.07, AME-ALE 0.04, PME-PME
0.14, PME-PLE 0.11, ALE-PLE 0.05.
Sternum 1.15 mm long and 0.95 mm
wide. Chelicerae 0.71 mm long with teeth
as in male.
Abdomen 3.02 mm long, 2.11 nun wide.
Epigastric furrow 0.85 mm from tracheal
spiracle, spiracle 0.90 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (nun)
and indices: I 1.42, 18; II 1.48, 18; III 0.99,
25; IV 1.58, 16. Ventral spination: tibiae
I, II 2-2-2', III 1-1-2, IV 2-1-2; metatarsi
I, II 2-2-0, III 2-0-2, IV 2-2-2.
Epigynum as in Figure 11, internal geni-
talia as in Figure 12.
Natural history. Mature males have been
taken from late September through early
February, mature females from mid-Octo-
ber through mid- April. Specimens have
been taken from Spanish moss, by sweep-
ing in bottomland pine and hardwood
forests, by sifting leaves and by Malaise
trap.
Distribution. Mid-eastern states from
Long Lsland south to North Carolina, wost
to southern Illinois, eastern Missouri and
northern Alabama ( Map 1 ) .
Anyphaena crebrispina Chamberlin
Map 1; Figures 3, 20, 21, 22
Aiufpliacna crebrispina Chamberlin, 1919, Pomona
Coll. J. Ent. Zool., 12: 10, pi. 4, fig. 4 {$).
Male holotype from Clareniont, California, in
MCZ, examined. Bryant, 1931, Psyche. 38: 113,
pi. 6, fig. 11, $. Roewer, 1954, Katalog der
Araneae, 2: 528. Bonnet, 1955, Bibliographia
Araneoruni, 2: 343.
Amjphaena zina Chamberlin, 1919, Pomona Coll.
J. Ent. Zool., 12:11, pi. 4, fig. 5(9). Female
holotype from Clareniont, California, in MCZ,
examined. Roewer, 1954, Katalog der Araneae,
2: 530. Bonnet, 1955, Bibliographia Araneorum,
2: 349. NEW SYNONYMY.
Diagnosis. Anyphaena crebrispina is the
most aberrant member of the celer group,
but is most closely related to A. dixiatm.
220 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
Plate 4
Figures 35, 38. Left palpi, retrolateral view. Figures 36, 37, 39-42. Epigyna, ventral view. Figures 43-46, 48,
49. Internal genitalia, dorsal view. Figure 47. Anyphaenid tracheae, diagrammatic. Figure 50. Clubionid
tracheae, diagrammatic. 35. Anyphaena cochise new species. 36, 43. Anyphaena marginalis (Banks). 37, 44.
Anyphaena rita new species. 38. Anyphaena arbida new species. 39, 45. Anyphaena autumna new species.
40,46. Anyphaena gibboides new species. 41,48. /Anyp/7aeA?a ca?a//na new species. 42,49. Anyphaena hespar
new species.
Spidkh I-'amily Anyphaexidak • I'latnick 221
Males of A. cre])ris-}nna may be readily dis-
tinguished by the greatly expanded base
of the RTA (Fig. 20). If' this speeies were
known solely from the female, it would be
impossible to place it in the celer group:
the epigynum, with its greatly expanded
sidepieces and its hick of an c>xternally visi-
ble midpiece, is totally unlike that of any
other species in this group ( Fig. 21 ) .
Male (Los Angeles Co., California).
Coloration as in AnypJiaena celer. Total
length 4.61 mm. Carapace 2.00 mm long,
1.57 mm wide, cephalic width 0.74 mm,
clypeus height 0.10 mm. Eyes: diameters
(mm): AME 0.07, ALE 0.10, PME 0.10,
PLE 0.10; anterior eye row 0.43 mm long,
recurved; posterior eye row 0.56 mm long,
prociuved; MOQ length 0.25 mm, front
width 0.18 mm, back width 0.29 mm; eye
interdistances (mm): AME-AME 0.04,
AME-ALE 0.02, PME-PME 0.09, PME-
PLE 0.09, ALE-PLE 0.04.
Sternum 1.10 mm long, 0.S8 mm wide.
Chelicerae 0.55 mm long with 5 promar-
ginal teeth and 8 retromarginal denticles.
Abdomen 2.65 mm long, 1.58 mm wide.
Epigastric furrow 0.79 mm from tracheal
spiracle, spiracle 0.68 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I L69,
13; II 1.51, 15; III 1.06, 22; IV 1.69, 14.
Ventral spination: tibiae I 2-2-2*, II 1-2-
2\ III 1-2-2, IV 2-2-2; metatarsi I, II 2-
2-0, III 2-0-2, IV 2-2-2. Modifications of
third leg as in A. celer.
Palpus as in Figures 3, 20.
Female (Los Angeles Co., California).
Coloration as in male of A. celer. Total
length 4.39 mm. Carapace 1.85 mm long,
1.37 mm wide, cephalic width 0.77 mm,
clypeus height 0.08 mm. Eyes: diameters
(mm): AME 0.07, ALE 0.09, PME 0.09,
PLE 0.09; anterior eye row 0.41 mm long,
recurved; posterior eye row 0.56 mm long,
procurved; MOQ length 0.23 mm, front
width 0.18 mm, back width 0.28 mm; eye
interdistances (mm): AME-AME 0.04,
AME-ALE 0.02, PME-PME 0.10, PME-
PLE 0.08, ALE-PLE 0.04.
Sternum 1.08 mm long, 0.86 mm wide.
Chelieerac> 0.64 mm long with 4 promar-
ginal teeth and 9 retromarginal denticles.
Abdomen 2.99 nun long, 1.98 mm wide.
Epigastric furrow 0.95 mm from tracheal
spiracle, spiracle 0.90 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 1.39, 15; II 1.31, 16; III 0.75,
28; IV 1.44, 14. Ventral spination: tibiae
I 2-2-2*, II 1-2-0, III 1-1-0, IV 1-1-2;
metatarsi as in male.
Epigynum as in Figure 21, internal geni-
talia as in Figure 22.
Natural history. Mature males have been
taken in November, mature females from
early December through late April. Speci-
mens have been taken by Berlese funnel
sampling of grape bark.
Distribution. Central and southern Cali-
fornia ( Map 1 ) .
Anyphaena dixiana (Chamberlin and
Woodbury), new combination
Map 1; Figures 4, 23, 24, 25
Gaijcnna dixiana Chamberlin and Woodbun',
1929, Proc. Biol. Soc. Washington, 42: 138,
pi. 1, fig. 3 ( 9 ). Female holotype from St.
Ceorge, Utah, in AMNH, examined. Roewer,
1954, Katalog der Araneae, 2: 540 ( G. dixima
[sic]). Bonnet, 1957, Bibliographia Araneorum,
2: 1977.
Anyphaena coloradensis Bryant, 1931, Psyche,
38: 112, pi. 6, figs. 9, 10, pi. 7, figs. .30, 33
{ $, 9 ). Male holotype, female allotNpc from
Boulder, Colorado, in MCZ, examined. Roewer,
1954, Katalog der Araneae, 2: 528. Bonnet,
1955, Bibliographia Araneorum, 2: 343. NEW
SYNONYMY.
Diafinosis. This distinctive species is
closest to AmjpJuiena crebrispina, but may
be quickly recognized by the spur borne
on the dorsal prong of the RTA of males
(Fig. 25) and the greatly reduced epigynal
midpiece of females (Fig. 23).
Male (Cochise Co., Arizona). Coloration
as in Anyphaena celer except that posterior
.spiimerets have dorsal surface sharply di-
vided into dark brown lateral and pale
orange median halves.
Total length 3.85 mm. Carapace 1.67
222 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
mm long, 1.44 mm wide, cephalic width
0.65 mm, clypeus height 0.07 mm. Eyes:
diameters (mm): AME 0.06, ALE 6.09,
PME 0.09, PLE 0.10; anterior eye row 0.39
mm long, recurved; posterior eye row 0.51
mm long, procurved; MOQ length 0.25
mm, front width 0.16 mm, back width 0.26
mm; eye interdistances (mm): AME-
AME 0.04, AME-ALE 0.03, PME-PME
0.08, PME-PLE 0.08, ALE-PLE 0.05.
Sternum 0.96 mm long, 0.76 mm wide.
Chelicerae 0.53 mm long with 4 promar-
ginal teeth and 6 reti'omarginal denticles.
Endites slightly invaginated at middle.
Abdomen 2.56 mm long, 1.49 mm wide.
Epigastric furrow 0.76 mm from tracheal
spiracle, spiracle 0.76 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 1.69,
11; II 1.37, 14; III 0.81, 28; IV 1.44, 16.
Ventral spination: tibiae I, II 2-2-2*, III,
IV 1-2-2; metatarsi I, II 2-2-0, III 2-0-2,
IV 2-2-2. Modifications of third leg as in
A. celer.
Palpus as in Figures 4, 25.
Female (Cochise Co., Arizona). Colora-
tion as in male. Total length 4.14 mm.
Carapace 2.03 mm long, 1.57 mm wide,
cephalic width 0.86 mm, clypeus height
0.09 mm. Eyes: diameters (mm): AME
0.05, ALE 0.08, PME 0.09, PLE 0.10; an-
terior eye row 0.43 mm long, recurved;
posterior eye row 0.60 mm long, procurved;
MOQ length 0.26 mm, front width 0.20
mm, back width 0.32 mm; eye interdis-
tances (mm): AME-AME 0.09, AME-
ALE 0.05, PME-PME 0.15, PME-PLE
0.09, ALE-PLE 0.07.
Sternum 1.15 mm long, 0.86 mm wide.
Chelicerae 0.71 mm long with 5 promar-
ginal teeth and 8 retromarginal denticles.
Abdomen 2.50 mm long, 1.69 mm wide.
Epigastric furrow 0.60 mm from tracheal
spiracle, spiracle 0.67 mm from base of
.spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 1.46, 16; II 1.33, 17; III 0.94,
24; IV 1.49, 17. Ventral spination as in
male.
Epigynum as in Figure 23, internal geni-
talia as in Figure 24.
Natural history. Mature males have been
taken from mid- August through mid-May,
mature females from late September
through late April. Specimens have been
taken from 5400 to 9000 feet (1650-2750
m), in yellow pine/ oak and montane for-
ests, in alfalfa, under dead agave and fre-
quently in houses.
Distribution. Northcentral Colorado south
to western Texas, west to southern Cali-
fornia (Map 1 ) .
Anyphaena judicata O. P.-Cambridge
Map 1; Figures 5, 26, 33, 34
Anijphaena iiidicata O. P. -Cambridge, 1896,
Biologia Central! Americana, Aran., 1: 203, pi
26, fig. 4 { S ). Male holotype from Omiltemi,
Guerrero, Mexico, in BMNH, examined. F. O.
P.-Cambridge, 1900, Biologia Centrali Ameri-
cana, Aran., 2: 96, pi. 7, fig. 9, $. Roewer,
1954, Katalog der Araneae, 2: 525. Bonnet,
1955, Bibliographia Araneormn, 2: 345.
Diagnosis. Anyphaena judicata is most
closely related to an unnamed Mexican
species (or group of species) and has no
close relatives among the species occur-
ring north of Mexico. Males may be easily
recognized by the distinctive form of the
dorsal prong of the RTA (Fig. 26). The
female epigynum is closest to that of A.
niarginalis, but the midpiece is proportion-
ately shorter and wider and the sidepieces
are narrower and diminish in width an-
teriorly smoothly, without the sharp de-
crease in width shown by A. marginalis
(Fig. 33).
Male (Cochise Co., Arizona). Coloration
as in Anyphaena celer, except that pos-
terior spinnerets have entire dorsal surface
dark brown.
Total length 3.46 mm. Carapace 1.76
mm long, 1.44 mm wide, cephalic width
0.68 mm, clypeus height 0.09 mm. Eyes:
diameters (mm): AME 0.06, ALE 0.10,
PME 0.09, PLE 0.10; anterior eye row 0.40
mm long, recurved; posterior eye row 0.52
mm long, procurved; MOQ length 0.26
mm, front width 0.17 mm, back width
Spider Family Anyphaenidae • Plotnick 223
0.28 mm; eye interdistanees (mm): AME-
AME 0.05, AME-ALE 0.03, PME-PME
0.11, PME-PLE 0.06, ALE-PLE 0.04.
Sternum 0.95 mm long, ().6S mm wide.
Chelicerae 0.56 mm long with 4 promar-
ginal teeth and 7 retromarginal denticles.
Abdomen 1.80 mm long, 1.15 mm wide.
Epigastric furrow 0.61 mm from tracheal
spiracle, spiracle 0.63 mm from base of
.spinnerets.
Tibial lengths (mm) and indices: I 2.25,
6; II 1.93, 8; III 1.01, 21; IV 1.66, 11. Ven-
tral spination: tibiae I 4-2-2*, II 3-2-2',
III 1-2-0, IV 1-1-2; metatarsi I, II, 2-2-0,
III 2-0-2; IV 1-2-2. Femur III unmodi-
fied. Tibia III ventral spine 1 on retrolat-
eral side missing, ventral spine 2 thickened,
cone-like. Coxae I, II and III ( but not IV )
with a small number of short, thick setae.
Coxae III with a tiibercule.
Palpus as in Figures 5, 26.
Female (Cochise Co., Arizona). Colora-
tion as in male.
Total length 4.72 mm. Carapace 1.76
mm long, 1.37 mm wide, cephalic width
0.81 mm, clypeus height 0.06 mm. Eyes:
diameters (mm): AME 0.07, ALE 0.10,
PME 0.10, PLE 0.10; anterior eye row 0.44
mm long, recurved; posterior eye row 0.60
mm long, procurved; MOQ length 0.29
mm, front width 0.21 mm, back width 0.32
mm; eye interdistanees (mm): AME-
AME 0.07, AME-ALE 0.03, PME-PME
0.13, PME-PLE 0.09, ALE-PLE 0.05.
Sternum 0.97 mm long, 0.77 mm wide.
Chelicerae 0.58 mm long with teeth as in
male.
Abdomen 3.13 mm long, 2.09 mm wide.
Epigastric furrow 1.21 mm from tracheal
spiracle, spiracle 1.31 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 1.69, 11; II 1.31, 14; III 0.88,
23; IV 1.66, 12. Ventral spination as in
male.
Epigynum as in Figure 33, internal geni-
talia as in Figure 34.
Natural history. Mature males have been
taken from mid-June through mid-August,
mature females from late March to Novem-
ber, most ill July and August. Specimens
have been taken from 5100 to 8000 feet
(1550-2450 m), by sweeping and under
rocks.
Distribution. Arizona south to Guerrero,
Mexico (Map 1).
Anyptiaena marginalis (Banks),
new combination
Map 1; Figures 6, 27, 36, 43
Gayeima marginalis Banks, 1901, Proc. Acad.
Natur. Sci. Philadelphia, 53: 574, pi. 2.3, fig.
22 ( 9 ). Female holotype from Beulali, San
Miguel Co., New Mexico, was probabl>' de-
posited in the MCZ along with the other types
from this paper but was not found by Bryant
when the MCZ t>pes were cataloged; lost,
presumed destroyed. Roewer, 1954, Katalog
der Araneae, 2: 540. Bonnet, 19.57, Biblio-
graphia Araneorum, 2: 1978.
Diagnosis. Amjphaena marginalis is most
closely related to A. hespar, both species
having a simple embolus and elongated
conductor. Males of A. marginalis (Fig.
27), however, do not have the spine-like
dorsal prong of the RTA of A. hespar, and
females of A. marginalis (Fig. 36) do not
have the conspicuous bulge in the epigynal
midpiece which characterizes A. hespar
females.
Male (Graham Co., Arizona). Colora-
tion as in Anyphaena celer.
Total length 3.78 mm. Carapace 1.98
mm long, 1.60 mm wide, cephalic width
0.72 mm, clypeus height 0.09 mm. Eyes:
diameters (mm): AME 0.06, ALE 0.10,
PME 0.08, PLE 0.10; anterior eye row 0.40
mm long, straight; posterior eye row 0.55
mm long, prociuved; xVlOQ length 0.20
mm, front width 0.17 mm, back width 0.28
mm; eye interdistanees (mm): AME-
AME 0.05, AME-ALE 0.02, PME-PME
0.11, PME-PLE 0.08, ALE-PLE 0.04.
Sternum 1.13 mm long, 0.81 mm wide.
Chelicerae 0.54 mm long with 5 promar-
ginal teeth and 6 retromarginal denticles.
Abdomen 2.00 mm long, 1.33 mm wide.
Epigastric furrow 0.52 mm from traclieal
224
Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
spiracle, spiracle 0.59 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 1.67,
13; II 1.35, 16; III 1.03, 26; IV 1.62, 14.
Ventral spination: tibiae I 4-2-2*, II 2-2-
2*, III, IV 1-2-2; metatarsi I, II 2-2-0, III
2-0-2, IV 2-2-2. Femur III unmodified.
Tibia III ventral spine 1 on retrolateral
side missing. Coxae unmodified.
Palpus as in Figures 6, 27.
Female (Graham Co., Arizona). Colora-
tion as in male of A. celer.
Total length 4.26 mm. Carapace 2.11
mm long, 1.55 mm wide, cephalic width
0.86 mm, clypeus height 0.08 mm. Eyes:
diameters (mm): AME 0.07, ALE 0.10,
PME 0.11, PLE 0.10; anterior eye row 0.44
mm long, straight; posterior eye row 0.64
mm long, procurved; MOQ length 0.30
mm, front width 0.19 mm, back width 0.33
mm; eye interdistances (mm): AME-
AME 0.05, AME-ALE 0.03, PME-PME
0.12, PME-PLE 0.09, ALE-PLE 0.06.
Sternum 1.05 mm long, 0.80 mm wide.
Chelicerae 0.65 mm long with 4 promar-
ginal teeth and 8 retromarginal denticles.
Abdomen 2.52 mm long, 1.53 mm wide.
Epigastric furrow 0.67 mm from tracheal
spiracle, spiracle 0.68 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 1.44, 18; II 1.21, 21; III 0.99,
25; IV 1.60, 17. Ventral spination as in
male except tibia III 1-1-2.
Epigynum as in Figure 36, internal geni-
talia as in Figure 43.
'Natural history. Mature males have been
taken from late August through late May,
mature females in all months except Janu-
ary and October. Specimens have been
taken from 6000 to 9300 feet (1850-2850
m), in yellow pine/ oak forests and under
rocks. I found this species in great abun-
dance by sorting pine litter at Rustler's
Park in the Chiricahua Mountains of south-
eastern Arizona in August 1972.
Distribution. Arizona, New Mexico and
Colorado (Map 1).
Anyphaena hespar new species
Map 1; Figures 16, 32, 42, 49
Types. Male holotype, female paratype
from Bear Canyon, Santa Catalina Moun-
tains, Pima Co., Arizona, 8 December 1968
(Karl Stephan), deposited in AMNH. Male
and female paratypes from Pima Co., Ari-
zona, deposited in MCZ. The specific
name is an arbitrary combination of letters.
Diagnosis. Anyphaena hespar is most
closely related to A. marginalis. Males of
the former may be distinguished by the
spine-like dorsal prong of their RTA (Fig.
32), females by the conspicuous bulge in
their epigynal midpiece (Fig. 42).
Male (Pima Co., Arizona). Coloration
as in Anyphaena celer.
Total length 3.13 mm. Carapace 1.62
mm long, 1.31 mm wide, cephalic width
0.59 mm, clypeus height 0.08 mm. Eyes:
diameters (mm): AME 0.05, ALE 0.08,
PxME 0.08, PLE 0.08; anterior eye row
0.33 mm long, straight; posterior eye row
0.45 mm long, procurved; MOQ length 0.19
mm, front width 0.14 mm, back width 0.24
mm; eye interdistances (mm): AME-
AME 0.05, AME-ALE 0.03, PME-PME
0.08, PME-PLE 0.07, ALE-PLE 0.04.
Sternum 0.95 mm long, 0.79 mm wide.
Chelicerae 0.39 mm long with 4 promar-
ginal teeth and 8 retromarginal denticles.
Abdomen 1.80 mm long, 1.10 mm wide.
Epigastric furrow 0.56 mm from tracheal
spiracle, spiracle 0.56 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 1.31,
17; II 1.08, 21; III 0.81, 28; IV 1.39, 18.
Ventral spination: tibiae I 4-2-2*, II 3-
2-2*, III 1-2-2, IV 2-2-2; metatarsi I, II
2-2-0, III 2-0-2, IV 2-2-2. Femur III un-
modified. Tibia III ventral spine 1 on retro-
lateral side missing, spine 2 thickened,
cone-like. Coxae unmodified.
Palpus as in Figures 16, 32.
Female (Pima Co., Arizona). Colora-
tion as in male of A. celer.
Total length 3.06 mm. Carapace 1.55
mm long, 1.26 mm wide, cephalic width
0.67 mm, clypeus height 0.06 mm. Eyes:
Spider Family Anvphaenidak • Plafuick 225
diameters (mm): AME 0.05, ALE O.OS,
PME 0.08, PLE O.OS; anterior eye row 0.33
mm long, .straight; posterior eye row 0.49
mm long, procurved; MOQ length 0.20
mm, front width 0.14 mm, back width 0.26
mm; eye interdistances (mm): AME-
AME 0.05, AME-ALE 0.03, PME-PME
0.10, PME-PLE 0.07, ALE-PLE 0.04.
Sternum 1.04 mm long, 0.70 mm wide.
Chelicerae 0.47 mm long with teeth as in
male.
Abdomen 1.85 mm long, 1.08 mm wide.
Epigastric furrow 0.49 mm from tracheal
spiracle; spiracle 0.41 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 1.13, 20; II 0.92, 25; III 0.72,
33; IV 1.26, 18. Ventral spination: tibiae
I, II 4-2-2^ III 1-1-2, IV 1-2-2; meta-
tarsi as in male.
Epigynum as in Figure 42, Internal geni-
talia as in Figure 49.
Natural history. Mature males and fe-
males have been taken from late October
through early April. Specimens have been
taken from leaf litter and under rocks.
DistriJmtion. Southeastern Arizona (Map
1)-
Anyphaena rita new species
Map 1; Figures 7, 28, 37, 44
Types. Male holotype, female paratype
from Bear Canyon, Santa Catalina Moun-
tains, Pima Co., Arizona, 8 December 1968
(Karl Stephan), deposited in AMNH. Male
and female paratypes from Pima Co., Ari-
zona, deposited in MCZ. The .specific
name is a noun in apposition derived from
the Santa Rita Mountains, where the
species is abundant.
Diagnosis. Anypliaemi rita is most closely
related to A. cochise, both species having
a conspicuously enlarged region of the
embolus and a slightly recurved tip of the
median apophysis. Males of A. rita (Fig.
28) may be distingui.shed by their smaller
size and by the differences in the dorsal
prong of the RTA. Females of A. cochise
are unknown, l)ut the epigynum of A. rita.
with its extremely broad midpiece, is quite
distinctive (Fig. 37).
Male (Pima Co., Arizona). Colorati(;n
as in Anyphaena celer.
Total length 4.10 mm. Carapace 1.94
mm long, 1.60 mm wide, cephalic width
0.67 mm, clypeus height 0.07 mm. Eyes:
diameters (mm): AME 0.05, ALE 0.08,
PME 0.09, PLE 0.09; anterior eye row 0.36
mm long, recurved; posterior eye row 0.53
mm long, procurved; MOQ length 0.22
mm, front width 0.15 mm, back width 0.27
mm; eye interdistances (mm): AME-
AME 0.05, AME-ALE 0.03, PME-PME
0.09, PME-PLE 0.09, ALE-PLE 0.05.
Sternum 1.13 mm long, 0.77 mm wide.
Chelicerae 0.50 mm long with 4 promar-
ginal teeth and 8 retromarginal denticles.
Abdomen 2.30 mm long, 1.26 mm wide.
Epigastric furrow 0.67 mm from tracheal
spiracle, spiracle 0.65 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 1.55,
14; II 1.39, 16; III 0.97, 23; IV 1.62, 14.
Ventral spination: tibiae I 4-2-2*, II
3-2-2*, III, IV 2-2-2; metatarsi I, II
2-2-0, III 2-0-2, IV 2-2-2. Femur III un-
modified. Tibia III ventral spines not
thickened. Coxae III and IV with only a
few short thick setae.
Palpus as in Figures 7, 28.
Female (Pima Co., Arizona). Coloration
as in male of A. celer.
Total length 5.04 mm. Carapace 2.05
mm long, 1.53 mm wide, cephalic width
1.03 mm, clypeus height 0.09 mm. Eyes:
diameters (mm): AME 0.05, ALE 6.10,
PME 0.10, PLE 0.11; anterior eye row 0.41
mm long, recurved; posterior eye row 0.58
mm long, procurved; MOQ length 0.32
mm, front width 0.18 mm, back width
0.29 mm; eye interdistances (mm): AME-
AME 0.07,' AME-ALE 0.03, PME-PME
0.09, PME-PLE 0.10, ALE-PLE 0.08.
Sternum 1.13 mm long, 0.81 mm wide.
Chelicerae 0.67 mm long with teeth as in
male.
Abdomen 2.75 mm long, 1.94 mm wide.
Epigastric furrow 1.06 mm from tracheal
226 Bulletin Museum of Comparative Zoologij, Vol. 146, No. 4
spiracle, spiracle 0.95 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 1.48, 15; II 1.26, 18; III
1.03, 21; IV 1.58, 17. Ventral spination as
in male except tibiae III, IV 1-2-2, meta-
tarsi IV 2-1-2.
Epigynum as in Figure 37, internal
genitalia as in Figure 44.
Natural history. Mature males have been
taken from mid-October through late
March, mature females from early June
tlirough early February. Specimens have
been taken from 4000 to 6800 feet. ( 1200-
2075 m), in oak/ grassland and under rocks.
Distribution. Arizona to Chihuahua,
Mexico (Map 1).
Anyphaena cochise new species
Map 1; Figures 13, 35
Types. Male holotype from Rustlers
Park, 8600 ft. (2625 m), Chiricahua Moun-
tains, Cochise Co., Arizona, 9 September
1950 (W. J. Gertsch), deposited in AMNH.
Male paratype from Cochise Co., Arizona,
deposited in MCZ. The specific name is a
noun in apposition and refers to the type
locality.
Diapiosis. Anypliaena cochise is most
closely related to A. vita, but the dorsal
prong of the RTA is relatively longer in
A. cochise (Fig. 35). Females of this
species are unknown.
Male (Cochise Co., Arizona). Colora-
tion as in Anyphaena celer.
Total length 5.44 mm. Carapace 2.52
mm long, 2.09 mm wide, cephalic width
0.88 mm, clypeus height 0.14 mm. Eyes:
diameters (mm): AME 0.09, ALE 6.13,
PME 0.13, PLE 0.13; anterior eye row
0.53 mm long, straight; posterior eye row
0.75 mm long, procurved; MOQ length
0.30 mm, front width 0.23 mm, back width
0.40 mm; eye interdistances (mm): AME-
AME 0.05, AME-ALE 0.04, PME-PME
0.14, PME-PLE 0.11, ALE-PLE 0.06.
Sternum 1.44 mm long, 1.08 mm wide.
Chelicerae 0.75 mm long with 4 promar-
ginal teeth and 7 retromarginal denticles.
Abdomen 3.38 mm long, 1.94 mm wide.
Epigastric fvuTow 0.92 mm from tracheal
spiracle, spiracle 1.03 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 2.32,
12; II 2.05, 13; III 1.39, 20; IV 2.14, 14.
Ventral spination: tibiae I 4-2-2*, II 2-2-
2*, III 1-2-2, IV 2-2-2; metatarsi I, II 2-
2-0, III 2-0-2, IV 2-2-2. Femur III un-
modified. Tibia III ventral spine 1 on ret-
rolateral side thickened slightly. All coxae
with a few scattered short thick setae.
Palpus as in Figures 13, 35.
Female. Unknown.
Natural history. Mature males have been
taken in early September at 8600 feet
(2625 m).
Distribution. Known only from the type
locality (Map 1).
Anyphaena autumna new species
IVIap 1; Figures 8, 29, 39, 45
Types. Male holotype, female paratype
from Rustler Camp, Chiricahua Mountains,
Cochise Co., Arizona, 9 September 1950
(W. J. Gertsch), deposited in AMNH.
Male and female paratypes from Cochise
and Graham Co., Arizona, deposited in
MCZ. The specific name refers to the
season of collection.
Diagno.sis. Anyphaena autumna is un-
likely to be confused with any other spe-
cies. The long recurved hook on the RTA
and the peculiar form of the tip of the
median apophysis are mil ike any other
species (Figs. 8, 29). The epigynum is
closest to that of A. gibboides, but the mid-
piece has a characteristic constriction near
its midpoint ( Fig. 39 ) .
Male (Cochise Co., Arizona). Colora-
tion as in Anyphaena celer, though the
paramedian bands on the carapace are
darker and wider than in that species.
Total length 5.51 mm. Carapace 2.50
mm long, 1.98 mm wide, cephalic width
1.03 mm, clypeus height 0.12 mm. Eyes:
Spider Family Anyphaenidae • Platnick 227
diameters (nini): AME 0.09, ALE 0.12,
PME 0.12, PLE 0.13; anterior eye row
0.55 mm long, recnrved; posterior eye row
0.75 mm long, proeun'ed; MOQ length
0.30 mm, front width 0.26 mm, back width
0.38 mm; eye interdistances (mm): AME-
AME 0.08, AME-ALE 0.05, PME-PME
0.15, PME-PLE 0.11, ALE-PLE 0.06.
Sternnm 1.46 mm long, 1.08 mm wide.
Chelicerae 0.79 mm long with 4 promar-
ginal teeth and 9 retromarginal denticles.
Abdomen 3.20 mm long, 2.16 mm wide.
Epigastric furrow 1.04 mm from tracheal
spiracle, spiracle 1.06 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 2.16,
13; II 1.93, 15; III 1.39, 22; IV 2.16, 14.
Ventral spination: tibiae I 2-2-2, II, III,
IV 1-2-2; metatarsi I, II 2-2-0, III 2-0-2,
I\^ 2-2-2. Third legs unmodified.
Palpus as in Figures 8, 29.
Female (Cochise Co., Arizona). Colora-
tion as in male.
Total length 6.41 mm. Carapace 2.34
mm long, 1.87 mm wide, cephalic width
1.12 mm, clypeus height 0.12 mm. Eyes:
diameters (mm): AME 0.10, ALE 0.13,
PME 0.13, PLE 0.13; anterior eye row
0.59 mm long, recurved; posterior eye row
0.70 mm long, procurved; MOQ length
0.33 mm, front width 0.27 mm, back width
0.42 mm; eye interdistances (mm): AME-
AME 0.06,' AME-ALE 0.03, PME-PME
0.17, PME-PLE 0.12, ALE-PLE 0.07.
Sternum 1.42 mm long, 1.08 mm wide.
Chelicerae 0.99 mm long with 4 promar-
ginal teeth and 8 retromarginal denticles.
Abdomen 3.96 mm long, 2.63 mm wide.
Epigastric furrow 1.33 mm from tracheal
spiracle, spiracle 1.33 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 1.75, 16; II 1.60, 18; III 1.10,
2.5; IV 1.89, 15. Ventral spination: tibiae
I 4-4-2, II 2-4-2, III 1-1-2, IV 1-2-2;
metatarsi as in male.
Epigynum as in Figure 39, internal geni-
talia as in Figure 45.
Natural history. Mature males and fe-
males have been taken in August and Sep-
tember. Specimens have be(Mi taken at
8200 fec>t (2500 m). I collected a few im-
matiue males (which matured in the labo-
ratory) of this .species in pine litter in the
Chiricahua Mountains, Arizona, where ma-
ture A. mar^inalis were extremely abun-
dant.
Dustribution. Southeastern Arizona (Map
1)-
Anyphaena gibboides new species
Map 1; Figures 14, 30, 40, 46
Types. Male holotype, female paratype
from City Creek Canyon, Salt Lake Co.,
Utah, 22 May 1943 (Wilton Ivie), depos-
ited in AMNH. Male and female para-
t)^es from Lake Co., Oregon, deposited in
MCZ. The specific name is an arbitrary
combination of letters.
Diagnosis. Anyphaena gihJ)oi(Ies is a
distinctive species. Males have a sharply
pointed median apophysis and serrate
RTA which will separate them from the
other known species (Figs. 14, 30). The
epigynum is closest to that of A. autumna,
but lacks the constriction of the midpiece
found in that species ( Fig. 40 ) .
Male (Salt Lake Co., Utah). Coloration
as in Anyphaena celer.
Total length 3.31 mm. Carapace 1.60
mm long, 1.28 mm wide, cephalic width
0.54 mm, clypeus height 0.07 mm. Eyes:
diameters (mm): AME 0.05, ALE 0.08,
PME 0.08, PLE 0.08; anterior eye row 0.34
mm long, straight; posterior eye row 0.48
mm long, procurved; MOQ length 0.23
mm, front width 0.15 mm, back width 0.24
mm; eye interdistances (mm): AME-
AME 0.05, AME-ALE 0.08, PME-PME
0.08, PME-PLE 0.07, ALE-PLE 0.05.
Sternum 0.85 mm long, 0.72 mm wide.
Chelicerae 0.49 mm long with 4 promar-
ginal teeth and 8 retromarginal denticles.
Abdomen 1.94 mm long, 1.24 mm wide.
Epigastric furrow 0.58 mm from tracheal
spiracle, spiracle 0.59 mm from base of
spinnerets.
228 BuUetm Museum of Comparative Zoology, Vol. 146, No. 4
Tibial lengths (mm) and indices: I 1.33,
17; II 1.24, 19; III 0.99, 23; IV 1.47, 16.
Ventral spination: tibiae I 2-2-0, II 1-2-0,
III 2-2-0, IV 2-2-2; metatarsi I, II 2-2-0,
III 2-1-2, IV 2-2-2. Modifications of third
leg as in A. celer save that all coxae have
clumps of short thick setae.
Palpus as in Figures 14, 30.
Female (Salt Lake Co., Utah). Colora-
tion as in male of A. celer.
Total length 3.74 mm. Carapace 1.75
mm long, 1.35 mm wide, cephalic width
0.83 mm, clypeus height 0.09 mm. Eyes:
diameters (mm): AME 0.06, ALE 0.09,
PME O.OS, PLE 0.08; anterior eye row
0.41 mm long, recurved; posterior eye row
0.57 mm long, procurved; MOQ length
0.24 mm, front width 0.18 mm, back width
0.28 mm; eye interdistances (mm): AME-
AME 0.06, AME-ALE 0.03, PME-PME
0.12, PME-PLE 0.09, ALE-PLE 0.06.
Sternum 1.19 mm long, 0.83 mm wide.
Chelicerae 0.62 mm long with 4 promar-
ginal teeth and 6 retromarginal denticles.
Abdomen 2.36 mm long, 1..39 mm wide.
Epigastric furrow 0.72 mm from tracheal
spiracle, spiracle 0.70 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 1.39, 18; II 1.24, 20; III
0.75, 27; IV 1.39, 18. Ventral spination:
tibiae I 2-2-0, II, III 1-2-0, IV 1-2-2;
metatarsi I, II 2-2-0, III 2-0-2, IV 2-2-2.
Epigynum as in Figure 40, internal geni-
talia as in Figure 46.
Natural history. Mature males and fe-
males have been taken in late May and
June. Habitat data is lacking.
Distribution. Northern Utah west to
southeastern Oregon ( Map 1 ) .
Anyphaena catalina new species
Map 1; Figures 15, 31, 41, 48
Types. Male holotype, female paratype
from Mt. Lemon, Santa Catalina Moun-
tains, Pima Co., Arizona, 13 July 1916 ( F.
E. Lutz), deposited in AMNH. Male and
female paratypes from Pima Co., Arizona,
and Mexico, Mexico, deposited in MCZ.
The specific name is a noun in apposition
and refers to the type locality.
Diagnosis. Anyphaena catalina is most
closely related to A. arbida, though males
of A. catalina may be readily distinguished
by their recurved retrolateral tegular
apophyses (Figs. 15, 31). Females of A.
arbida are unknown; those of A. catalina
may be recognized by the epigynal hood
being roughly equal in size to the epigynal
midpiece ( Fig. 41 ) .
Male (Pima Co., Arizona). Coloration
as in Anyphena celer.
Total" length 3.53 mm. Carapace 1.78
mm long, 1.42 mm wide, cephalic width
0.72 mm, clypeus height 0.09 mm. Eyes:
diameters (mm): AME 0.05, ALE 0.09,
PME 0.08, PLE 0.09; anterior eye row
0.40 mm long, recurved; posterior eye row
0.51 mm long, procurved; MOQ length
0.21 mm, front width 0.17 mm, back width
0.26 mm; eye interdistances (mm): AME-
AME 0.07, AME-ALE 0.04, PME-PME
0.09, PME-PLE 0.08, ALE-PLE 0.04.
Sternum 0.90 mm long, 0.70 mm wide.
Chelicerae 0.56 mm long with 4 promar-
ginal teeth and 6 retromarginal denticles.
Abdomen 1.85 mm long, 0.90 mm wide.
Epigastric furrow 0.61 mm from tracheal
spiracle, spiracle 0.65 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 2.07,
8; II 1.94, 9; III 1.08, 23; IV 1.80, 10. Ven-
tral spination: tibiae I 4-2-2*, II 2-2-2*,
III, IV 2-2-2; metatarsi I, II 2-2-0, III 2-
0-2, IV 2-2-2. Modifications of third leg
as in A. celer save that femur III lacks short
thick setae and all coxae bear clumps of
them.
Palpus as in Figures 15, 31.
Female (Pima Co., Arizona). Coloration
as in male of A. celer.
Total length 4.57 mm. Carapace 1.84
mm long, 1.42 mm wide, cephalic width
0.94 mm, clypeus height 0.09 mm. Eyes:
diameters (mm): AME 0.07, ALE 0.09,
PME 0.09, PLE 0.09; anterior eye row
0.47 mm long, recui-ved; posterior eye row
Spideu pAisriLY Anyphaemdak • Pintnick 229
0.63 mm long, prociirved; MOQ length 0.26
mm, front width 0.22 mm, back widtli 0.33
mm; eve interdi.stances (mm): AME-
AME 6.0S, AME-ALE 0.04, PME-PME
0.15, PME-PLE 0.11, ALE-PLE 0.07.
Sternum 1.01 mm long, 0.85 mm wide.
Chelicerae 0.68 mm long with 4 promar-
ginal teeth and 8 retromarginal denticle.s.
Abdomen 2.74 nnn long, 1.85 mm wide.
Epigastric furrow 0.86 mm from tracheal
.spiracle, spiracle 0.94 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 1.51, 14; II 1.33, 17; III 0.94,
23; IV 1.48, 16. Ventral spination: tibiae I,
II 2-2-2*, III 1-2-2, IV 2-2-2; metatarsi
as in male.
Epigynum as in Figure 41, internal geni-
talia as in Figure 48.
Natural Justory. Mature males and fe-
males have been taken in July and August.
Specimens have been taken at 7500 feet
(2300 m) in yellow pine/ oak and douglas
fir/ white fir forests.
Distribution. Southeastern Arizona south
to central Mexico (Map 1).
Anyphaena arbida new species
Map 1; Figures 17, 38
Types. Male holotype from Carr Can-
yon, Huachuca Mountains, Cochise Co.,
Arizona, 26 August 1950 (M. A. Cazier),
deposited in AMNH. Male paratype from
Cochise Co., Arizona, deposited in MCZ.
The specific name is an arbitrary combina-
tion of letters.
Diagnosis. AnypJiaena arhida is most
closely related to A. catalina. Males of the
former (Figs. 17, 38) lack the recurved
retrolateral tegular apophysis of A. cata-
lina; females of A. arbida are unknown.
Male (Cochise Co., Arizona). Colora-
tion as in AnypJiaena celer, except that
posterior spinnerets are as in A. dixiana.
Total length 6.95 mm. Carapace 3.28
mm long, 2.41 mm wide, cephalic width
1.22 mm, clypeus height 0.14 mm. Eyes:
diameter (mm): AME 0.11, ALE 0.13,
PME 0.13, PLE 0.15; anterior eye row
0.64 mm long, recurved; posterior eye row
0.89 mm long, procurved; MOQ lengtii
0.43 mm, front width 0.31 mm, back width
0.44 mm; eye interdistances (mm): AME-
AME 0.09, AME-ALE 0.04, PME-PME
0.18, PME-PLE 0.14, ALE-PLE 0.09.
Sternum 1.62 mm long, 1.33 mm wide.
Chelicerae 1.30 mm long with 4 promar-
ginal teeth and 8 retromarginal denticles.
Abdomen 3.71 mm long, 2.16 mm wide.
Epigastric furrow 1.08 mm from tracheal
spiracle, .spiracle 1.12 mm from base of
spinnerets. Spinnerets surrounded by a
clump of unusually long setae.
Tibial lengths (mm) and indices: I 6.88,
5; II 3.35, 10; III 2.20, 16; IV 3.35, 10.
Ventral spination: tibiae I 4-2-2*, II 3-
2-2*, III, IV 2-2-0; metatarsi I, II 2-2-0,
III, IV 2-2-2. Third legs unmodified.
Palpus as in Figures 17, 38.
Female. Unknown.
Natural history. Mature males have been
collected in August. Habitat data is lack-
ing-
1).
Di.stribution. Cochise Co., Arizona (Map
Anyphaena pectorosa Group
Diagnosis. The pectorosa group is closely
related to the pacifica group, but males
may be distinguished by the spins on their
coxae (Figs. 59-62). Females have the
epigynum on a characteristic sclerotized
plate (Figs. 74, 77, 79) and simple .sper-
mathecae (Figs. 75, 78, 80).
Description. Total length 4.5-6.5 mm.
Carapace longer than wide, narrowed in
front to less than half its maximum width
in males, to slightly more than half its
maximum width in females. Clypeus height
more than 1.5 times the diameter of an an-
terior median eye. Posterior median, pos-
terior lateral and anterior lateral eyes sub-
equal in size, almost twice the diameter of
anterior medians. Procurved posterior eye
row longer than slightly recur\xKl anterior
row. Median ocular (juadrangle almost
230 Bulletin Museum of Comparative Zoolofi.ij, Vol. 146, No. 4
twice as wide in back as in front. Anterior
median eyes separated l:)y sliglitly less than
their diameter, sHghtly closer to anterior
laterals than to each other. Posterior me-
dians separated by slightly more than their
diameter, slightly closer to posterior lat-
erals. Anterior laterals separated by their
radins from posterior laterals. Sternum
longer than wide, with a low hirsute knob
behind its middle in some males. Chelic-
erae with 4 promarginal teeth and 7-9
retromarginal denticles. Abdomen longer
than wide, tiacheal spiracle midway be-
tween epigastric furrow and base of spin-
nerets. Leg formula 1423. Metatarsi I
and 11 with two pairs of ventral spines.
Males with coxae II bearing round knobs,
coxae III and IV bearing spurs. Palpus
with an elongated median apophysis, en-
larged conductor and inconspicuous embo-
lus. Retrolateral tibial apophysis short.
Epigynum on a sclerotized plate, without
a hood. Two simple spermathecae.
Variation. The species in this group
show little intraspecific variation, individ-
ual or geographical, in size, structure or
coloration.
Key to Species
la. Coxae III of males with posterior spur bifid
(Fiffs. 59, 61, 62); sternum of males with a
low hirsute knob behind middle; sclerotized
epigynal plate wider posteriorly than an-
teriorly (Figs. 74, 79) 2
lb. Coxae III of males with posterior spur not
bifid ( Fig. 60 ) ; sternum of males without a
low hirsute knob behind middle; sclerotized
epigynal plate wider anteriorly than pos-
teriorly (Fig. 77) fratema
2a. Distal tip of palpal median apophysis bent
sharply towards cymbium (Figs. 55, 58);
sclerotized epigynal plate with pronounced
posterolateral corners ( Fig. 74 ) 3
2b. Distal tip of palpal median apophysis not
bent sharply towards cymbium ( Fig.
57 ) ; sclerotized epigynal plate without pro-
nounced posterolateral corners (Fig. 79)
alaclma
3a. Distal tip of palpal median apophysis meet-
ing the recessed, dorsal branch of tlie apoph-
ysis ( Fig. 55 ) ; sclerotized epigynal plate
with pronounced posterolateral comers
( Fig. 74 ) pectorosa
3b. Distal tip of palpal median apophysis not
meeting the recessed, dorsal branch of the
apophysis (Fig. 58); females unknown __7flc?:a
Anyphaena pectorosa L. Koch
Map 2; Figures 51, 55, 59, 74, 75
Anyphaena pectorosa L. Koch, 1866, Arachn. Fam.
Drass., 198, pi. 8, figs. 131, 132 { $). Male
holotype from Baltimore, Maryland, in BMNH,
examined. Bryant, 1931, Psyche, 38: 110, pi. 6,
fig. 5, $ . Chickering, 1939, Pap. Michigan
Acad. Sci., 24: 51, figs. 5-8, $,9. Comstock,
1940, Spider Book, rev. ed., p. 577, fig. 636, 9 .
Kaston, 1948, Bull. Connecticut Geol. Natur.
Hist. Surv., 70: 408, figs. 1453, 1477-1480,
$, 9. Roewer, 1954, Katalog der Araneae, 2:
529. Bonnet, 1955, Bibliographia Araneorum,
2: 346.
Aniiphaena calcarata Emerton, 1890, Trans. Con-
necticut Acad. Sci., 8: 187, pi. 6, figs. 3-3d ( $,
9 ). Male holotype, female allotype from West
Haven, Connecticut, in MCZ, examined. Emer-
ton, 1902, Common Spiders, p. 12, figs. 42, 43,
$, 9.
Gaijenna calcarata, Banks, 1910, Bull. U.S. Nat.
Mus., 72: 13.
Gaijenna pectorosa, Comstock, 1912, Spider Book,
p. 563 (in part), fig. 636, 9 (not fig. 637).
Diagnosis. Anyphaena pectorosa is closest
to A. alachua, but may readily be distin-
guished from it by the highly curved me-
dian apophysis of males (Fig. 55) and the
pronounced posterolateral corners of the
sclerotized epigvnal plate of females (Fig.
74).
Male (Fairfax Co., Virginia). Total
length 5.40 mm. Carapace 2.43 mm long,
1.98 mm wide, cephalic width 0.88 mm,
clypeus height 0.11 mm, yellow with thin
dark border and two dark paramedian
longitudinal bands. Eyes: diameters (mm):
Plate 5
Figures 51--54. Left palpi, ventral view. Figures 55-58. Left palpi, retrolateral view. Figures 59-62.
ventral view. 51, 55, 59. Anyptiaena pectorosa L. Koch. 52, 56, 60. Anyphaena fraterna (Banks).
Anyphaena alachua new species. 54, 58, 62. Anyphaena lacka new species.
Male coxae,
53, 57, 61.
Spider I'^aafily Anyphaenidak • Plaliuck 231
232 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
AME 0.06, ALE 0.11, PME 0.11, PLE 0.12;
anterior e\e row 0.48 mm long, slightly re-
cur^•ed; posterior eye row 0.65 mm long,
procurved; MOQ length 0.28 mm, front
width 0.20 mm, back width 0.35 mm; eye
interdistances (mm): AME-AME 0.07,
AME-ALE 0.04, PME-PME 0.14, PME-
PLE 0.13, ALE-PLE 0.05.
Sternum 1.35 mm long, 1.01 mm wide,
pale yellow with translucent border, dark-
ened extensions to coxae and a low hirsute
knob behind middle. Chelicerae 0.73 mm
long with 4 promarginal teeth and 7 retro-
marginal denticles, pale yellow with boss
outlined in gray. Labium and endites yel-
low, darkest proximally. Endites slightly
invaginated at middle.
Abdomen 3.15 mm long, 1.67 mm wide,
pale white with transverse rows of dark
markings, venter pale. Epigastric furrow
1.01 mm from tracheal spiracle, spiracle
1.06 mm from base of spinnerets.
Legs pale yellow with distal segments
darkest. Tibial lengths (mm) and indices:
I 3.10, 7; II 2.52, 9; III 1.82, 16; IV 2.56, 10.
Ventral spination: tibiae I 2-2-1, II-IV 2-
2-2; metatarsi I, II 2-2-0, III 2-0-2, IV 2-
2-2. Coxae II, III and IV modified as in
Figure 59.
Palpus as in Figures 51, 55.
Female (Fairfax Co., Virginia). Colora-
tion as in male.
Total length 5.44 mm. Carapace 2.41
mm long, 1.91 mm wide, cephalic width
0.97 mm, clypeus height 0.08 mm. Eyes:
diameters (mm): AME 0.07, ALE 0.12,
PME 0.11, PLE 0.12; anterior eye row
0.52 mm long, recurved; posterior eye row
0.71 mm long, procurved; MOQ length
0.33 mm, front width 0.20 mm, back width
0.37 mm; eye interdistances (mm): AME-
AME 0.05, AME-ALE 0.04, PME-PME
0.15, PME-PLE 0.10, ALE-PLE 0.07.
Sternum 1.31 mm long, 1.06 mm wide,
without hirsute knob. Chelicerae 0.72 mm
long with 4 promarginal teeth and 8 retro-
marginal denticles.
Abdomen 3.10 mm long, 1.76 mm wide.
Epigastric furrow 0.70 mm from tracheal
spiracle, spiracle 1.22 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 2.41, 11; II 2.05, 13; III 1.44,
19; IV 2.20, 12. Ventral spination: tibiae
I, II 2-2-0, III, IV 1-2-1; metatarsi I, II
2-2-0, III, IV 2-2-2.
Epigynum as in Figure 74, internal geni-
talia as in Figure 75.
Natural history. Mature males have been
taken from mid-April through early Sep-
tember, mature females from mid-April
through mid-August. Specimens have been
taken by sweeping foliage, in Malaise and
pitfall ti^ips, and under rocks. Egg cases
taken with females contained 65-95 eggs.
Distribution. New England west to
Michigan, south to western Florida and
eastern Texas ( Map 2 ) .
Anyphaena atachua new species
Map 2; Figures 53, 57, 61, 79, 80
Types. Male holotype, female paratype
from west of Gainesville, Alachua Co.,
Florida, 18 April 1938 (Willis J. Certsch),
deposited in AMNH. Male and female
paratypes from Alachua Co., Florida, de-
posited in MCZ. The specific name is a
noun in apposition and refers to the type
locality.
Diagnosis. Anyphaena alachua is closest
to A. pectorosa but the median apophysis
is not highly curved (Fig. 57) and the
epigynal plate lacks pronounced postero-
lateral corners (Fig. 79).
Male (Alachua Co., Florida). Colora-
tion as in Anyphaena pectorosa.
Total length 4.90 mm. Carapace 2.41
mm long, 2.01 mm wide, cephalic width
0.79 mm, clypeus height 0.13 mm. Eyes:
diameters (mm): AME 0.07, ALE 0.12,
PME 0.12, PLE 0.13; anterior eye row
0.51 mm long, slightly recurved; posterior
eye row 0.70 mm long, procurved; MOQ
length 0.30 mm, front width 0.22 mm, back
width 0.36 mm; eye interdistances (mm):
AME-AME 0.07, AME-ALE 0.04, PME-
PME 0.12, PME-PLE 0.11, ALE-PLE 0.06.
Spider Family Anvi'haemdae • Plalnick 233
Sternum 1.26 mm long, 1.01 mm wide,
with low hirsute knob Ix^hind middle.
Chelicerae 0.76 mm long with 4 promar-
ginal teeth r.nd 9 retromarginal denticle.s.
Abdomen 2.48 mm long, 1.48 mm wide.
Epigastric furrow 0.76 mm from tracheal
spiracle, spiracle 0.8.3 mm from base of
spinnerc>ts.
Tibial lengtlis (nun) and indices: I 2.77,
10; II 2.27,^11; III 1.44, 22; IV 1.94. 14.
\Vntral spination: tibiae I, II 2-2-0, III
1-2-2, IV 2-2-2; metatarsi I, II 2-2-0, III
2-0-2, IV 2-2-2. Coxae II, III and I\'
modified as in Figure 61.
Palpus as in Figures 53, 57.
Female (Alachua Co., Florida). Colora-
tion as in male of A. pectorosa.
Total length 6.17 mm. Carapace 2.45
mm long, 1.80 mm wide, cephalic width
0.94 mm, clypeus height 0.12 mm. Eyes:
diameters (mm): AME 0.08, ALE 0.13,
PME 0.12, PLE 0.13; anterior eye row
0.57 mm long, slightly recurved; posterior
eye row 0.73 mm long, procui-ved; MOQ
IcMigth 0.30 mm, front width 0.22 mm, back
width 0.40 mm; eye interdistances (mm):
AME-AME 0.07, AME-ALE 0.04, PME-
PME 0.15, PME-PLE 0.11, ALE-PLE
0.07.
Sternum 1.35 mm long, 1.08 mm wide,
without hirsute knob. Clielicerae 0.84 mm
long with teeth as in male.
Abdomen 3.53 mm long, 2.02 mm wide.
Epigastric furrow 1.10 mm from tracheal
spiracle, spiracle 1.21 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 2.30, 13; II 1.91, 14; III 1.31,
22; IV 2.09, 13. Ventral spination as in
male save metatarsi III 2-2-2.
Epigynum as in Figure 79, internal geni-
talia as in Figure 80.
Natural history. Mature males have been
taken in late April and early May, mature
females from late March through mid-May,
by sweeping.
Distribution. Known only from Florida
(Map 2).
Anyphaena lacka new species
Map 2; Figures 54, 58, 62
Type. Male liolotxpe from Lake Corpus
Christi State Park, southwest of Mathis,
San Patricio Co., Texas, 28 Jvme 1962 (J. A.
Beatty), deposited in MCZ. The specific
name is an arbitrary combination of letters.
Dia<i,nosis. Anyphaena lacka is most
closely reflated to A. alachua ])ut has a dis-
tinct point on the tip of the median apoph-
ysis (Fig. 58). Females of A. lacka are
unknown.
Male (San Patricio Co., Texas). Colora-
tion as in Anyphaena pectorosa.
Total length 4.61 mm. Carapace 2.05
mm long, 1.69 mm wide, cephalic width
0.79 mm, clypeus height 0.12 mm. Eyes:
diameters (mm): AME 0.07, ALE 0.12,
PME 0.11, PLE 0.11; anterior eye row
0.47 mm long, slightly recurved; posterior
eye row 0.61 mm long, procurved; MOQ
length 0.26 mm, front width 0.19 mm, back
width 0.32 mm; eye interdistances (mm):
AME-AME 0.05, AME-ALE 0.03, PME-
PME 0.11, PME-PLE 0.08, ALE-PLE 0.04.
Sternum 1.24 mm long, 0.90 mm wide,
with low hirsute knob behind middle.
Chelicerae 0.64 mm long with 4 promar-
ginal teeth and 9 retromarginal denticles.
Abdomen 2.41 mm long, 1.33 mm wide.
Epigastric furrow 0.74 mm from tracheal
spiracle, spiracle 0.90 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 2.38,
9; II 1.91, 12; III 1.32, 30; IV 1.93, 12.
Ventral spination: tibiae I, II 2-2-0, III,
IV 1-2-2; metatarsi I, II 2-2-0, III 2-0-2,
IV 2-2-2. Coxae II, III, and W modified
as in Figure 62.
Palpus as in Figures 54, 58.
Female. LTnknown.
Natural history and distribution. Known
only from the type specimen.
Anyphaena fraterna (Banks)
iVlap 2; Figures 52, 56, 60, 77, 78
.\]iil})hacna coiispcrsa KeyserlinK. 1887, Verh.
zool. bot. Cos. Wien, 37: 453, pi. 6, fig. 23
234 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
( $ ). Female holotype from Bee Spring, Ken-
tucky, in MCZ, examined; preoccupied by Any-
phaena conspersa Simon, 1878.
Gaijenna fratema Banks, 1896, Trans. Amer. Ent.
Soc, 23: 63. Male holotype from Sea Cliff,
New York, in MCZ, examined.
Amjphaena fratema, Simon, 1897, Hist. Natur.
Araign., 2: 96. Bryant, 1931, Psyche, 38: 110,
pi. 6, fig. 6, pi. 8, fig. 23, $, 9. Comstock,
1940, Spider Book, rev. ed., p. 577, fig. 637, $ .
Kaston, 1948, Bull. Connecticut Geol. Natur.
Hist. Surv., 70: 408, figs. 1454-1456, $, 9.
Roewer, 1954, Katalog der Araneae, 2: 529.
Bonnet, 1955, Bibliographia Araneonnn, 2: 344.
Sillus consperstis, Petrunkevitch, 1911, Bull. Amer.
Mus. Natur. Hist., 29: 511.
Gayenna pectorosa, Comstock, 1912, Spider Book,
p. 563 (in part), fig. 637, $.
Diagnosis. Anyphaena fratema is a dis-
tinctive species easily recognized by the
long and narrow median apophysis of
males (Fig. 52) and by the female's epigy-
nal plate being wider anteriorly than pos-
teriorly (Fig. 77).
Male (Hall Co., Georgia). Coloration
as in Anyphaena pectorosa.
Total length 4.93 mm. Carapace 2.23
mm long, 1.85 mm wide, cephalic width
0.81 mm, clypeus height 0.10 mm. Eyes:
diameters (mm): AME 0.06, ALE 0.12,
PME 0.11, PLE 0.11; anterior eye row 0.45
mm long, slightly recurved; posterior eye
row 0.64 mm long, procurved; MOQ length
0.30 mm, front width 0.18 mm, back width
0.33 mm; eye interdistances (mm): AME-
AME 0.05, AME-ALE 0.04, FME-PiME
0.12, PME-PLE 0.08, ALE-PLE 0.05.
Sternum 1.28 mm long, 0.99 mm wide,
without hirsute knob. Chelicerae 0.59 mm
long with 4 promarginal teeth and 9 retro-
marginal denticles.
Abdomen 2.83 mm long, 1.60 mm wide.
Epigastric furrow 0.97 mm from tracheal
spiracle, spiracle 0.85 mm from base of
spinnerets.
Legs with scattered dark spots. Tibial
lengths (mm) and indices: I 2.60, 8; II
2.16, 11; III 1.52, 16; IV 2.47, 10. Ventral
.spination: tibiae I, II 2-2-0, III 1-2-2, IV
2-2-2; metatarsi I, II 2-2-0, III, IV 2-2-2.
Coxae II, III and IV modified as in Figure
60.
Palpus as in Figures 52, 56.
Female (Hall Co., Georgia). Coloration
as in male of A. pectorosa.
Total length 5.00 mm. Carapace 2.32
mm long, 1.80 mm wide, cephalic width
0.94 mm, clypeus height 0.10 mm. Eyes:
diameters (mm): AME 0.06, ALE 0.11,
PME 0.11, PLE 0.11; anterior eye row
0.49 mm long, recurved; posterior eye row
0.69 mm long, procui-ved; MOQ length
0.27 mm, front width 0.18 mm, back width
0.36 mm; eye interdistances (mm): AME-
AME 0.05,' AME-ALE 0.04, PME-PME
0.15, PME-PLE 0.09, ALE-PLE 0.05.
Sternum 1.28 mm long, 1.04 mm wide.
Chelicerae 0.75 mm long with teeth as in
male.
Abdomen 2.97 mm long, 1.71 mm wide.
Epigastric furrow 0.85 mm from tracheal
spiracle, spiracle 0.85 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 2.29, 11; II 1.89, 13; III 1.30,
19; IV 2.16, 13. Ventral spination as in
male.
Epigynum as in Figure 77, internal geni-
talia as in Figure 78.
Natural history. Mature males have been
taken from late March through early July,
mature females from late March through
late August. Specimens have been taken
by sweeping foliage, in Malaise and pitfall
traps, and under logs. I collected this spe-
cies in great abundance by sweeping
honeysuckle (Lonicera sp.) at night in
southern West Virginia during June 1971.
Distribution. Southern New York west
to eastern Kansas, south to western Florida
and eastern Texas (Map 2).
Anyphaena pacifica Group
Diagnosis. The pacifica group is closely
related to the pectorosa group and appears
to displace it in the western United States.
The males have similarly short retrolateral
tibial apophyses (Figs. 69-71), but paci-
fica group males lack the coxal spurs char-
acteristic of the pectorosa group, though
Spider Family ANYPiiAiixiDAt; • Flatnick 235
<:
Anyphoena pectorosa o\ -* >
I
------v*-
i£
Anyphoena fraterna .«\ -^>
f* ■'
1
c "^
1
ijiy. \
'(
VM • ^
1
r^"^- *'
1
1
)• ;' ''*~
• .-••\
•
_
(• '~~^-r- ;
• 1
V ;'•
_ 1
~'^
";•:• z---
..^
• • -
i
r^^
1
Anyphaenc
pacificQ
/ A
-^w
Tn P
^■-^ .__..__
Anyphoena
colifornicQ -
1
\\v
1
1 "~
r )
Anyphoena
locko
s , 1
\\
i ,'---,'
; .' t>'-
^. ^ ■'
■• "> /
\
\ 1
"\ : 1
Anyphoena aperta '
\ 1 -^ ^ .'-
Map 2. Distributions of Anyphaena alachua, A. aperta, A. californica, A. fraterna, A. gertschi, A. lacka, A. pacif-
ica and A. pectorosa.
males of Anyphaena gertschi have rounded
knobs on the coxae. Females lack the
sclerotized epigynal plates found in the
pectorosa group, but have a lightly sclero-
tized atrivun-like area posteromedially
(Figs. 66, 67, 72) and long, sometimes
coiling, ducts (Figs. 68, 73, 76).
Description. Total length 4-6 mm. Cara-
pace longer than wide, narrowed in front
by at least one-third of its maximum width,
often by more than half. Clypeus height
roughly equal to anterior median eye diam-
eter. All eyes subequal in size. Procurved
posterior eye row longer than slightly re-
curved anterior eye row. Median ocular
quadrangle longer than wide in front,
wider in back than long. Anterior median
eyes separated by less than their diameter,
much closer to anterior laterals than to
each other. Posterior medians separated
by more than their diameter, much closer
to posterior laterals. Anterior laterals sepa-
rated by slightly more than their radius
from posterior laterals. Sternum longer
than wide, without a hirsute knob. Chelic-
erae with 3 promarginal teeth and 6-9
retromarginal denticles. Abdomen longer
than wide, tracheal spiracle midway be-
tween epigastric furrow and base of spin-
nerets. Leg formula 1423. Metatarsi I and
II with two pairs of ventral spines. Males
with legs unmodified. (A. pacifica and A.
californica) or with coxae bearing round
knobs and femora II and III bearing
patches of short stiff setae ventrally (A.
gertschi). Palpus with an elongated me-
dian apophysis, enlarged conductor and
inconspicuous embolus. Retrolateral tibial
apophysis short. Epigyiuim not on a scle-
rotized plate, without a hood, with a more
or less pronoimced atrium-like lightly
sclerotized area posteromedially. Internal
genitalia with long ducts that coil in some
species.
Variation. Two species in this group, A.
pacifica and A. californica, show a great
deal of xariation in genitalic structure. In
both species the shape oi the tip of the pal-
236 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
pal median apophysis and the coihng of
the epigynal ducts are strikingly variable,
and it was initially thought that many
species were involved. Three sources of
evidence, however, have indicated other-
wise. First, many females are found in
which the ducts on one side of the epigy-
num coil differently from those on the
other side. Secondly, when many speci-
mens are taken together at one locality on
a single da}", several variants are often
found. Finally, the retrolateral tibial
apophysis, which usually provides excellent
diagnostic characters in anyphaenids, is
stable within the species as they are de-
fined here. Until such time as biological
evidence on the breeding habits of these
spiders can be obtained, it seems best to
consider both A. pacifica and A. califorjuco
as widespread, variable species.
Key to Species
la. Retrolateral tibial apophysis (RTA) without
a dorsal process (Fig. 69). Median apoph-
ysis with a deep invagination below tip
giving the tip a chelate appearance (Fig.
65). Epigyninii with large wing-shaped
paramedian flaps ( Fig. 72 ) gertschi
lb. Retrolateral tibial apophysis (RTA) with a
dorsal process (Figs. 70, 71). Median
apophysis without a deep invagination be-
low tip (Figs. 63, 64). Epigynum with-
out large wing-shaped paramedian flaps
(Figs. 66, 67) 2
2a. Dorsal process of RTA short, located dis-
tally (Fig. 70). Median apophysis narrow-
ing gradually towards tip (Fig. 63). In-
ternal ducts with many coils (Fig. 68) — _
— pacifica
2b. Dorsal process of RTA long, located prox-
inially (Fig. 71). Median apophysis nar-
rowing abruptly towards tip (Fig. 64). In-
ternal ducts without many coils (Fig. 73)
— californica
Anyphaena pacifica (Banks)
Map 2; Figures 63, 66, 68, 70
Gaijcnna pacifica Banks, 1896, Trans. Amer. Ent.
Soc, 23: 63. Female holot>'pe from Olympia,
Washington, in MCZ, examined.
Anyphaena pacifica, Simon, 1897, Hist. Natur.
Araign., 2: 96. Bryant, 1931, Psyche, 38: 115,
pi. 8, fig. 36, ?. Levi and Levi, 1951, Zoo-
logica (New York), 36: 228, Tig. 25, $. Roe-
wer, 1954, Katalog der Araneae, 2: 529. Bon-
net, 1955, Bibliographia Araneorum, 2: 346.
Anyphaena mundella Chamberlin, 1920, Pomona
Coll. J. Ent. Zool., 12: 12, pi. 5, fig. 3 ( 9 , not
$, = Aysha incursa) . Female holotype from
Claremont, California, in MCZ, examined.
Bryant, 1931, Psyche, 38: 120 (sub Aysha de-
cepta [sic] ). Roewer, 1954, Katalog der Araneae
2: 534 (sub Aysha decepta [sic]). Bonnet,
1955, Bibliographia Araneorum, 2: 836 (sub
Aysha decepta [sic]). NEW SYNONYMY.
Anyphaena intermontana Chamberlin, 1920,
Canad. Ent., 52: 200, fig. 22-6 ( $ ). Female
holotype from Mill Creek, Salt Lake Co., Utali,
in MCZ, examined. Bryant, 1931, Psyche, 38:
114 (sub Anyphaena californica [sic]). Roe-
wer, 1954, Katalog der Araneae, 2: 528 (sub
Anyphaena californica [sic]). Bonnet, 1955,
Bibliographia Araneorum, 2: 343 (sub Any-
phaena californica [sic]). NEW SYNONYMY.
Gayenna saniuana Chamberlin and Gertsch, 1928,
Proc. Biol. Soc. Wash., 41: 185. Male holotype
from Verdure, San Juan Co., Utah, in AMNH,
examined. Roewer, 1954, Katalog der Araneae,
2: 540. NEW SYNONYMY.
Anyphaena saniuana, Bryant, 1931, Psyche, 38:
107. Bonnet, 1955, Bibliographia Araneorum,
2: 347.
Anyphaena pomona Chamberlin and Ivie, 1941,
Bull. Univ. Utah, Biol., 6: 23, pi. 2, fig. 16
( 9 ). Female holotype from Mill Creek, Te-
hama Co., California, in AMNH, examined.
Roewer, 1954, Katalog der Araneae, 2: 529.
NEW SYNONYMY.
Gayenna jollensis Schenkel, 1950, Verb. Naturf.
Ges. Basel, 61: 77, fig. 27 ( 9 ). Female holo-
type from La Jolla, California, in Naturhistor-
isches Museum, Basel, examined. Roewer, 1954,
Katalog der Araneae, 2: 540. NEW SYN-
ONYMY.
Plate 6
Figures 63-65. Left palpi, ventral view. Figures 69-71. Left palpal tibiae, retrolateral view. Figures 66, 67, 72,
74, 77, 79. Epigyna, ventral view. Figures 68, 73, 75, 76, 78, 80. Internal genitalia, dorsal view. 63, 66, 68, 70.
Anyptiaena pacifica {Banks). 64,67,71,73. Anyptiaena californica {Banks). 65,69,72,76. Anyphaena gertschi
new species. 74, 75. Anyphaena pectorosa L. Koch. 77, 78. Anyphaena fraterna (Banks). 79, 80. Anyphaena
alachua new species.
Spider Family Anyphaenidae • Plalnick 237
68
^V
?)
78
•^
£S^'
238 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
Diagnosis. Amjphaena pacifica is closest
to A. californica, but males may be distin-
guished by the short, distal, dorsal process
of the retrolateral tibial apophysis (Fig.
70) and the gradually narrowing tip of the
median apophysis (Fig. 63), while females
have distinctive highly coiled internal
ducts (Fig. 68). Variation in this species
is discussed above.
Male (El Dorado Co., California). Total
length 5.18 mm. Carapace 2.34 mm long,
1.94 mm wide, cephalic width 0.86 mm,
clypeus height 0.12 mm, pale orange with
thin dark border and two dark paramedian
longitudinal bands. Eyes: diameters (mm):
AME 0.09, ALE 0.12, PME 0.10, PLE 0.11;
anterior eye row 0.51 mm long, slightly
procurved; posterior eye row 0.69 mm long,
procurved; MOQ length 0.28 mm, front
width 0.24 mm, back width 0.34 mm; eye
interdistances (mm): AME-AME 0.07,
AME-ALE 0.03, PME-PME 0.14, PME-
PLE 0.10, ALE-PLE 0.05.
Sternum 1.49 mm long, 1.04 mm wide,
pale orange with darker border. Chelicerae
0.67 mm long with 3 promarginal teeth
and 8 retromarginal denticles, dark orange-
brown proximally, pale orange distally,
with boss outlined in gray. Labium and
endites orange, darkest proximally. En-
dites slightly invaginated at middle.
Abdomen 2.81 mm long, 1.69 mm wide,
reddish-brown throughout. Epigastric fur-
row 0.85 mm from tracheal spiracle, spira-
cle 0.92 mm from base of spinnerets.
Legs pale orange, unmodified. Tibial
lengths (mm) and indices: I 2.11, 12; II
1.87, 13; III 1.44, 20; IV 2.07, 15. Ventral
spination: tibiae I, II 2-2-0, III 1-2-2, IV
2-2-2; metatarsi I, II 2-2-0, III, IV 2-2-2.
Palpus as in Figures 63, 70.
Female (Mono Co., California). Color-
ation as in male.
Total length 5.39 mm. Carapace 2.34
mm long, 1.62 mm wide, cephalic width
0.94 mm, clypeus height 0.09 mm. Eyes:
diameters (mm): AME 0.10, ALE 0.12,
PME 0.11, PLE 0.11; anterior eye row 0.51
mm long, slightly recurved; posterior eye
row 0.73 mm long, procurved; MOQ length
0.29 mm, front width 0.25 mm, back width
0.36 mm; eye interdistances (mm): AME-
AME 0.06, AME-ALE 0.03, PME-PME
0.15, PME-PLE 0.10, ALE-PLE 0.07.
Sternum 1.44 mm long, 1.01 mm wide.
Chelicerae 0.71 mm long with teeth as in
male.
Abdomen 3.02 mm long, 1.69 mm wide.
Epigastric furrow 0.81 mm from tracheal
spiracle, spiracle 0.86 mm from base of
spinerets.
Tibial lengths (mm) and indices: I 1.84,
15; II 1.71, 15; III 1.39, 19; IV 2.07, 13.
Ventral spination as in male save tibiae
III 1-1-2 and IV 1-2-2.
Epigynum as in Figure 66, internal geni-
talia as in Figure 68.
Natural history. Mature males have been
taken from late February through late
July, mature females year round. Speci-
mens have been taken in montane forests,
in pitfall traps, under rocks and commonly
in houses.
Distribution. Western North America
from British Columbia south to California,
Ai-izona and New Mexico (Map 2).
Anyphaena californica (Banks)
Map 2; Figures 64, 67, 71, 73
Gaijenna californica Banks, 1904, Proc. California
Acad. Sci., 3: 338, pi. 38, fig. 2(9). Female
holotype from Palo Alto, California, in MCZ,
examined.
Amiphaena mens Chamberlin, 1920, Pomona Coll.
J. Ent. Zoo!., 12: 11, pi. 5, fig. \ {$). Male
holotype from Claremont, California, in MCZ,
examined. Bryant, 1931, Psyche, 38: 113.
Roewer, 1954, Katalog der Araneae, 2: 529.
Bonnet, 1955, Bibliographia Araneorum, 2: 347.
NEW SYNONYMY.
Anyphaena californica, Bryant, 1931, Psyche, 38:
114. Roewer, 1954, Katalog der Araneae, 2:
528. Bonnet, 1955, Bibliographia Araneorum,
2: 343.
Diagnosis. Amjphaena californica is most
closely related to A. pacifica, but males
have a long, proximal, dorsal process on
the retrolateral tibial apophysis (Fig. 71)
and an abruptly narrowed tip of the me-
Spider Family Anyphaenidae • Phifnick
239
diaii apopliysis (Fig. 64), while the inter-
nal dncts of the female are not highly
coiled (Fig. 73). Variation in this species
is discnssed above.
Male (San Diego Co., California). Col-
oration as in AnypJiaena pacifica except
that the abdomen is pale white with trans-
verse rows of dark markings.
Total length 4.68 mm. Carapace 2.21
mm long, 1.78 mm wide, cephalic width
0.68 mm, clypeus height 0.07 mm. Eyes:
diameters (mm): AME 0.07, ALE 6.09,
PME 0.10, PLE 0.11; anterior eye row
0.43 mm long, recm-ved; posterior eye row
0.59 mm long, procurved; MOQ length
0.30 mm, front width 0.20 mm, back width
0.32 mm; eye interdistances (mm): AME-
AME 0.06, AME-ALE 0.04, PME-PME
0.13, PME-PLE 0.11, ALE-PLE 0.07.
Sternum 1.31 mm long, 0.90 mm wide.
Chelicerae 0.60 mm long with 3 promar-
ginal teeth and 8 retromarginal denticles.
Abdomen 2.97 mm long, 1.34 mm wide.
Epigastric furrow 0.79 mm from tracheal
spiracle, spiracle 0.85 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 3.28,
5; II 3.20, 7; III 2.27, 8; IV 2.93, 7. Ventral
spination: tibiae I 2-2-0, II 2-2-2, III 1-
1-2, IV 1-2-2; metatarsi I, II 2-2-0, III,
I\' 2-2-2.
Palpus as in Figures 64, 71.
Female (Humboldt Co., CaHfornia).
Coloration as in male.
Total length 5.98 mm. Carapace 2.56
mm long, 1.91 mm wide, cephalic width
1.03 mm, clypeus height 0.08 mm. Eyes:
diameters (mm): AME 0.09, ALE 0.11,
PME 0.12, PLE 0.12; anterior eye row 0.52
mm long, recurved; posterior eye row 0.69
mm long, procurved; MOQ length 0.35
mm, front width 0.25 mm, back width 0.37
mm; eye interdistances (mm): AME-
AME 0.07, AME-ALE 0.04, PME-PME
0.13, PME-PLE 0.10, ALE-PLE 0.06.
Sternum 1.44 mm long, 1.08 mm wide.
Chelicerae 0.86 mm long with 3 promar-
ginal teeth and 9 retromarginal denticles.
Abdomen 3.64 mm long, 2.43 mm wide.
Epigastric furrow 1.15 mm from tracheal
spiracle, spiracle 1.33 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 2.16,
12; II 1.87, 13; III 1.30, 19; IV 2.06, 14.
Ventral spination as in male except tibiae
II 1-2-0.
Epigynum as in Figure 67, internal geni-
talia as in Figure 73.
Natural Jii.story. Mature males have been
taken from early March through mid-July,
mature females from mid-March through
mid-November. Specimens have Ix^en
taken in redwood forests, on citrus trees
and in houses.
Distribution. Oregon and California
(Map 2).
Anyphaena gertschi new species
Map 2; Figures 65, 69, 72, 76
Types. Male holotype, female paratype
from Bluff, San Juan Co., Utah, 11 May
1933 (Wilton Ivie), deposited in AMNH.
Male and female paratypes from Emery
Co., Utah, deposited in MCZ. The specific
name is a patronym in honor of Willis J.
Gertsch, who first recognized the species
as new.
Diagnosis. Anyphaena gertschi is a dis-
tinctive species easily recognized by the
chelate appearance of the tip of the me-
dian apophysis of males (Fig. 65) and
the large wing-shaped paramedian flaps
on the female epigynum (Fig. 72).
Male (Emery Co., Utah). Coloration as
in Anyphaena pacifica except that cara-
pace has paramedian bands only vaguely
indicated and abdomen is pale yellow
throughout.
Total length 4.00 mm. Carapace 1.85
mm long, 1.42 mm wide, cephalic width
0.92 mm, clypeus height 0.14 mm. Eyes:
diameters (mm): AME 0.09, ALE 6.09,
PME 0.09, PLE 0.09; anterior eye row
0.45 mm long, slightly recurv'ed; pos-
terior eye row 0.59 mm long, procurved;
MOQ length 0.26 mm, front width 0.22
mm, back width 0.32 mm; e\e interdis-
240 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
tances (mm): AME-AME 0.04, AME-
ALE 0.03, PME-PME 0.14, PME-PLE
0.08, ALE-PLE 0.04.
Sternum 1.12 mm long, 0.85 mm wide.
Chelicerae 0.65 mm long with 3 promar-
ginal teeth and 6 retromarginal denticles.
Abdomen 2.11 mm long, 1.31 mm wide.
Epigasti'ic furrow 0.67 mm from tracheal
spiracle, spiracle 0.76 mm from base of
spinnerets.
All coxae with round knobs ventrally.
Femora II and III with patches of short,
thick setae ventrally. Tibial lengths (mm)
and indices: I 2.00, 9; II 1.69, 13; III 1.30,
17; IV 1.87, 12. Ventral spination: tibiae
I 2-2-0, II 1-2-0, III, IV 1-2-2; metatarsi
I, II 2-2-0, III, IV 2-2-2.
Palpus as in Figures 65, 69.
Female (San Diego Co., Cahfornia).
Coloration as in male.
Total length 5.04 mm. Carapace 2.25 mm
long, 1.76 mm wide, cephalic width 0.95
mm, clypeus height 0.12 mm. Eyes: diam-
eters (mm): AME 0.10, ALE 0.13, PME
0.10, PLE 0.13; anterior eye row 0.51 mm
long, straight; posterior eye row 0.68 mm
long, procurved; MOQ length 0.28 mm,
front width 0.26 mm, back width 0.36 mm;
eye interdistances (mm): AME-AME
0.06, AME-ALE 0.03, PME-PME 0.16,
PME-PLE 0.08, ALE-PLE 0.05.
Sternum 1.28 mm long, 0.90 mm wide.
Chelicerae 0.70 mm long with teeth as in
male.
Abdomen 3.10 mm long, 2.02 mm wide.
Epigastric furrow 0.77 mm from tracheal
spiracle, spiracle 1.03 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 1.62, 14; II 1.49, 15; III 1.17,
21; IV 1.69, 14. Ventral spination as in
male save tibiae III 1-1-0.
Epigynum as in Figure 72, internal geni-
talia as in Figure 76.
Natural history. Mature males have been
taken from late April through late June,
mature females from mid-May through
late September. Nothing is known of the
habits of this species.
Distribution. Southern Utah south to
southern California and Arizona (Map 2).
Anyphaena accentuata Group
Diagnosis. Members of this group can
be immediately differentiated from the
other nearctic Anyphaena by the presence
of only one pair of ventral spines on meta-
tarsi I and II. Only one species occurs in
America north of Mexico.
Description. Total length 4-6 mm. Cara-
pace longer than wide, narrowed in front
to less than half its maximum width in
males, to slightly more than half in females.
Clypeus height roughly equal to anterior
median eye diameter. Median eyes smaller
than laterals. Procurved posterior eye row
longer than recurved anterior row. Me-
dian ocular quadrangle longer than wide
in front, wider in back than long. Anterior
median eyes separated by less than their
diameter, closer to anterior laterals. Pos-
terior medians separated by 1.5 times their
diameter, closer to posterior laterals. An-
terior laterals separated by their radius
from posterior laterals. Sternum longer
than wide, unmodified. Chelicerae with 3
promarginal teeth and 5-7 retrolateral den-
ticles. Abdomen longer than wide, tracheal
spiracle midway between epigastric furrow
and base of spinnerets. Leg formula 1423,
legs unmodified. Metatarsi I and II with
one pair of ventral spines. Palpus with
short median apophysis, short conductor
and conspicuous embolus. Cymbial groove
compressed to retrolateral side of cymbium.
Epigynum with hood. Internal genitalia
with anterior membranous dorsal cover.
Variation. No significant variation was
detected in Anyphaena aperta.
Anyphaena accentuata (Walckenaer)
Figure 134
Aranea accentuata Walckenaer, 1802, Faun. Paris,
2: 226. Type lost, presumed destroyed.
Anyphaena accentuata, Roewer, 1954, Katalog der
Araneae, 2: 522. Bonnet, 1955, Bibliographia
Araneoriini, 2: 338.
Spider Family Axypiiaenidak • Plafnick 241
A drawing of tlic palpus of tliis Enro-
pc^an spider, type species ol tlie genus A/ij/-
pJiaena, is included for pinposes of com-
parison to A. aperta. Confusion exists
between AnypJiaena accentuatii, A. ohscura
(Sundex'all) and A. sabina L. Koch, and
the female is therefore not illustrated and
no description is gi\^en. The male illus-
trated is from England.
Anyphaena aperta (Banks)
Map 2; Figures 135-137
Gaijenna aperta Banks, 1921, Pioc. California
Acad. Sci., 11: 100, fig. ,3 ( 9 ). Female holo-
t\'pe from OKinpia, Washington, in MCZ, ex-
amined.
Ani/pliacna aperta, Bryant, 1931, Psyche, 38: 114,
pi. 8, fig. 35, 9 . Fox, 1938, Iowa State Coll. J.
Sci., 12: 238, pi. 1, fig. 6, $. Roewer, 1954,
Katalog der Araneae 2: 528. Bonnet, 1955,
Bibliographia Araneornm, 2: 342.
Didiinosis. In addition to the diagnostic
character of the species group, Amjphaemi
aperta can readily be distinguished from
all other North American anyphaenids by
the sharply pointed median apophysis of
males (Fig. 135) and the membranous dor-
sal cover of the internal genitalia of females
(Fig. 137). Although the distribution indi-
cates that this might be an inti-oduced
species, no specimens or described species
from the Palearctic or Oriental regions re-
semble Anyphaena aperta.
Male (Yamhill Co., Oregon). Total
length 4.32 mm. Carapace 1.98 mm long,
1.63 mm wide; cephalic width 0.74 mm,
clypeus height O.OS mm, light orange-
brown, darker towards sides, with two
dark paramedian longitudinal bands. Eyes:
diameters (mm): AME 0.07, ALE 0.11,
PME 0.09, PLE 0.11; anterior eye row
0.44 mm long, recurved; posterior eye row
0.62 mm long, procurved; MOQ length
0.26 mm, front width 0.20 mm, back width
0.32 mm; eye interdistances (mm): AME-
AME 0.05, AME-ALE 0.03, PME-PME
0.14, PME-PLE 0.10, ALE-PLE 0.06.
Sternum 1.04 mm long, 0.89 mm wide,
pale orange \\'ith translucent border and
darkened extensions to coxae. Chelicerae
0.55 mm long with 3 promarginal teeth and
5 retromarginal denticles, orange-brown
with boss outlined in gray. Labium and
endites pale orange, darkest proximally.
Endites not invaginated.
Abdomen 2.52 mm long, 1.51 mm wide,
pale white with transx'crse rows of dark
markings, venter pale with a clump of
thick elongate setae posteriorly. Epigastric
furrow 0.86 mm from tracheal spiracle,
spiracle 0.74 mm from base of spinnerets.
Legs pale yellow, unmodified. Tibial
lengths (mm) and indices: I 1.87, 12; II
1.70, 13; III 1.27, 18; IV 1.73, 14. Ventral
spination: tibiae I, II 2-2-2, III 1-2-2,
IV 2-2-2; metatarsi I, II 2-0-0, III 2-0-2,
IV 2-2-2.
Palpus as in Figure 135.
Female (Curry Co., Oregon). Colora-
tion as in male.
Total length 5.83 mm. Carapace 2.65
mm long, 2.05 mm wide, cephalic width
1.17 mm, clypeus height 0.09 mm. Eyes:
diameters (mm): AME 0.10, ALE 0.12,
PME 0.12, PLE 0.13; anterior eye row
0.61 mm long, slightly recurved; posterior
eye row 0.87 mm long, procurved; MOQ
length 0.35 mm, front width 0.30 mm, back
width 0.44 mm; eye interdistances (mm):
AME-AME 0.09, AME-ALE 0.05, PME-
PME 0.19, PME-PLE 0.14, ALE-PLE 0.07.
Sternum 1.46 mm long, 1.04 mm wide.
Chelicerae 0.80 mm long with 3 promar-
ginal teeth and 7 retromarginal denticles.
Abdomen 4.00 mm long, 2.60 mm wide,
without thick setae ventrally. Epigastric
furrow 0.81 mm from tracheal spiracle,
spiracle 1.03 mm from base of spinnerets.
Tibial lengths (mm) and indices: I 1.87,
16; II 1.77, 16; III 1.31, 22; IV 1.87, 17.
Ventral spination as in male except tibiae
I 2-2-0 and IV 1-2-2.
Epigynum as in Figure 136, internal
genitalia as in Figure 137.
Natural history. Mature males have been
taken from late Marcli through early Sep-
tember, mature females from ],\tc March
242 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
through early November. Specimens have
been taken from redwoods and red cedars.
Distribution. Pacific coast from British
Coknnbia south to southern Cahfornia
(Map 2).
Wulfila O. P.-Cambridge
WuIfiJa O. P.-Cambridge, 1895, Biologia Central!
Americana, Aran., 1: 158. Type species Wulfila
pallidtis O. P.-Cambridge, 1895, designated by
Simon, 1897, Hist. Natur. Araign., 2: 103.
Cragits O. P.-Cambridge, 1896, Biologia Centrali
Americana, Aran., 1: 215. Type species by mono-
typy Cragiis pallidus O. P.-Cambridge, 1896.
NEW SYNONYMY.
Anyphaenella Bryant, 1931, Psyche, 38: 115. Type
species by original designation Clubiona salta-
hunda Hentz, 1847. NEW SYNONYMY.
Diagnosis. Wulfila may be easily recog-
nized by their long, thin, pale white legs.
Leg I in particular is greatly elongated,
with its tibial index usually 5 or less. Pal-
pal structure indicates that this genus is
closely related to Amjphaena. There are
probably more than fifty species in this
genus; most occur in Central America and
the West Indies.
Description. Total length 2.5-4.5 mm.
Carapace longer than wide, narrowed in
front to from one-half to two-thirds its
maximum width. Clypeus height greater
than anterior median eye diameter. Pos-
terior median, posterior lateral and anterior
lateral eyes subequal in size, somewhat
larger than anterior medians. Procurved
posterior eye row longer than straight an-
terior row. Median ocular quadrangle
twice as wide in back as in front. Anterior
median eyes separated by less than their
diameter, by roughly their diameter from
anterior laterals. Posterior medians sepa-
rated by almost twice their diameter, by
their diameter from posterior laterals. An-
terior laterals separated by roughly their
diameter from posterior laterals. Sternum
longer than wide, unmodified. Chelicerae
with 3-6 promarginal teeth, often on ca-
rina, and 5-10 retromarginal denticles.
Abdomen longer than wide, tracheal spira-
cle midway between epigastric furrow and
base of spinnerets. Leg formula 1423, legs
long, thin, pale white. Leg I greatly elon-
gated. Metatarsi I and II with two pairs
of ventral spines. Coxae of males often
with spurs and knobs; leg III spination
often reduced. Palpus with an elongated
median apophysis, enlarged conductor and
conspicuous embolus. Retrolateral tibial
apophysis greatly expanded except in W.
wunda. Epigyna and internal genitalia
small and diverse.
Variation. None of the species in this
genus show any significant individual or
geographic intraspecific variation in struc-
ture, size or coloration.
Key to Species
la. Carapace and abdomen with dark mark-
ings saltabunda
lb. Carapace and abdomen without dark
markings 2
2a. Males 3
2b. Females . 7
3a. At least one pair of coxae modified with
spurs or knobs 4
3b. All coxae unmodified alba
4a. Coxae I and/or II modified with spurs or
knobs 5
4b. Coxae III and/or IV modified with spurs
or knobs - 6
5a. Retrolateral tibial apophysis more than
half the tibial length (Fig. 93) — bryantae
5b. Retrolateral tibial apophysis less than half
the tibial length ( Fig. 95 ) ._ wunda
6a. Retrolateral tibial apophysis greatly ex-
panded at tip (Fig. 86) ..._. tantilh
6b. Retrolateral tibial apophysis not greatly
expanded at tip (Fig. 88) immaculella
7a. Epigynum with long ducts (Figs. 91, 97,
98 ) 8
7b. Epigynum without long ducts (Figs. 90,
96) : 10
8a. Epigynum with a heart-shaped atrium
(Fig. 97) wunda
8b. Epigynum without a heart-shaped atrium 9
9a. Epigynal ducts terminating far anterior of
epigynal openings ( Fig. 91 ) tantilla
9b. Epigynal ducts temiinating near epigynal
openings ( Fig. 98 ) immaculella
10a. Epigynum with anterolateral flaps, with-
out a medial ridge (Fig. 90) alba
10b. Epigynum without anterolateral flaps, with
a medial ridge (Fig. 96) bryantae
Spideh Family Anyphaenidaf, • Platnick 243
^;
V)
i
)
!-•-
\U^
•^
•,
rf ?
•
^ _- •^-^•.^%
•\
\/^
Wulfila saltabundQ^
1
--—— ^^"^
^^-j
^
\
Ifila tantilla
1
V
i
r
i
1
1
■^
Wulfila immaculella |
VV
\
U '^
— u-^/- ...
r
Wulfllo albo
\-- -
1 ' ■
1 — -'i
V .'
N
.
^"^ /^
v-4
/■•f
\
\
Wulfllo bryantae
"\
Map 3. Distributions of Teudis calcar, Wulfila alba, W. bryantae, W. immaculella, W. saltabunda, W. tantilla and
IV. wunda.
Wulfila pallidas O. P.-Cambridge
Figure 144
Wulfila palUdus O. P.-Cambridge, 1895, Biologia
Central! Americana, Aran., 1: 159, pi. 19, fig.
11 ( 9 ). Female holotype from Teapa, Ta-
basco, Mexico, in BMNH, examined. Bonnet,
1959, Bibliographia Araneorum, 2: 4832.
Wulfila pallida, Simon, 1897, Hist. Natur. Araign.,
2: 94. Roewer, 1954, Katalog der Araneae, 2:
554.
Vulfila pallida, Simon, 1897, Hist. Natur. Araign.,
2: 103.
Thi.s Mexican .species, though belonging
to a distinct species group, closely resem-
bles the North American Wulfila in body
form, leg length and coloration. It is the
type .species of Wulfila.
Wulfila saltabunda (Hentz),
new combination
Map 3; Figures 81, 82, 89, 99
Cluhiona saltabunda Hentz, 1847, J. Boston Soc.
Natur. Hist., 5: 453, pi. 23, fig. 23 ( 9 ). Fe-
male holotype from Alabama in Boston Soc.
Natur. Hist. (Boston Mu.seum of Science), de-
stroyed by beetles.
Amjphaena saltabunda, Fmerton, 1890, Trans.
Connecticut Acad. Sci., 8: 187, figs. 4-4d, $,
$ . Emerton, 1902, Common Spiders, p. 14,
figs. 46, 47, 5 , 9 .
Gayenna saltabunda, Comstock, 1912, Spider
Book, p. 563, figs. 638, 639, $, 9 .
Anyphaenella saltabunda, Bryant, 1931, Psyche,
38: 116, pi. 7, figs. 18, 22, $, 9. Comstock,
1940, Spider Book, rev. ed., p. 576, figs. 638,
639, $, 9. Ka.ston, 1948, Bull. Connecticut
Geol. Natur. Hist. Surv., 70: 406, figs. 1465-
1470, $, 9. Roewer, 1954, Katalog der
Araneae, 2: 530. Bonnet, 1955, Bibliographia
Araneorum, 2: 349.
Dia<i,nosis. Wulfila saltabunda is the
only species in this area which has dark
markings on the carapace and abdomen.
In addition, the shape of the retrolateral
tibial apophysis ( Fig. 82 ) and sperma-
thecae (Fig. 99) serve to distinguish it
from W. alba, its closest relative.
Male (Suffolk Co., New York). Total
length 3.06 mm. Carapace 1.46 mm long,
1.04 mm wide, cephalic width 0.54 mm,
clypeus height 0.07 mm, pale white with
thin dark border and two dark paramedian
longitudinal bands. Eyes: diameters
(mm): AME 0.05, ALE 0.09, PME 0.09,
PLE 0.09; anterior eye row 0.39 mm long,
straight; posterior eye row 0.50 mm long,
244 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
86
^■'^.,
■■ 90
Plate 7
Figures 81, 83, 85, 87. Left palpi, ventral view. Figures 82, 84, 86, 88. Left palpi, retrolateral view. Figures
89-91. Epigyna, ventral view. 81. 82, 89. Wulfila saltabunda (Hentz). 83, 84, 90. Wulfila alba (Hentz). 85, 86,
91. Wulfila tantilla Chickering. 87, 88. Wulfila immaculella (Gertsch).
Spider Faaiily Anyphaenidae • Phifnick
245
procurx'ed; MOQ length 0.22 mm, front
width 0.14 mm, back width 0.30 mm; eye
interdistances (mm): AME-AME 0.04,
AME-ALE 0.03, PME-PME 0.12, PiME-
PLE 0.06, ALE-PLE 0.03.
Sternum 0.86 mm long, 0.59 mm wide,
pale white with thick tran.slucent border
witli extensions to coxae and large triangu-
lar dark spots between coxae. Chelicerae
0.40 mm long with 6 promarginal teeth and
7 retromarginal denticles, pale white with
boss outlined in gray and several very long
setae. Laliium and endites pale white,
endites not in\'aginated at middle.
Abdomen 1.60 mm long, 0.97 mm wide,
pale white with transverse rows of dark
spots, venter with thin dark median line
anterior of epigastric furrow and two large
median dark spots between epigastric fur-
row and spinnerets. Epigastric furrow 0.40
mm from tracheal spiracle, spiracle 0.45
nmi from base of spinnerets.
Legs pale white, unmodified, though leg
III spination reduced. Tibial length (mm)
and indices: I 2.70, 4; II 1.42, 9; III 1.08,
13; IV 1.55, 10. Ventral spination: tibiae
I 2-2-0, II 1-1-0, III 0-1-0, IV 1-1-0;
metatarsi I, II 2-2-0, III 0-0-0, IV 1-2-2.
Palpus as in Figures 81, 82.
Female (Suffolk Co., New York). Col-
oration as in male.
Total length 4.18 mm. Carapace 1.78
mm long, 1.28 mm wide, cephalic width
0.70 mm, clypeus height 0.09 mm. Eyes:
diameters (mm): AME 0.06, ALE 0.09,
PME 0.09, PLE 0.10; anterior eye row 0.43
mm long, straight; posterior eye row 0.58
mm long, procurved. MOQ length 0.28
mm, front width 0.15 mm, back width 0.32
mm; eye interdistances (mm): AME-
AME 0.04, AME-ALE 0.05, PME-PME
0.14, PME-PLE 0.08, ALE-PLE 0.04.
Sternum 0.99 mm long, 0.74 mm wide.
Chelicerae 0.56 mm long with 6 promar-
ginal teeth and 10 retromarginal denticles.
Abdomen 2.47 mm long, 2.27 mm wide.
Epigastric furrow 0.85 mm from tracheal
spiracle, .spiracle 0.85 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 2.56,
5; II 1.31, 12; III 0.90, 20; IV 1.67, 12.
\'entral spination as in male except tibiae
II 2-2-0 and III 1-1-0 and metatarsi III
2-1-0.
Epigynum as in Figure 89, internal geni-
talia as in Figure 99.
Natural Jiistonj. Mature males have been
taken from mid-April through late August,
mature females from late April through
late August. Specimens have been taken
by sweeping and on apple trees.
Distrilnifion. Nova Scotia west to Min-
nesota and Nebraska, south to Florida and
eastern Texas ( Map 3 ) .
Wulfila alba (Hentz),
new combination
Map 3; Figures 83, 84, 90, 100
Cluhiona albeus Hentz, 1847, J. Boston See.
Natur. Hist., 5: 454, pi. 23, fig. 24 {$). Male
holotype from Alabama in Boston Soc. Natur.
Hist. (Boston Museum of Science), destroyed
by beetles.
Anyphaetia alhcns, Marx, 1883, in Howard, A List
of the Invertebrate Fauna of South Carolina, p.
24.
Chimcanthium alhens, Mar.x, 1890, Proc. U.S. Nat.
Mus., 12: 513.
AmiphaeucUa alba, Bryant, 1931, Psyche, 38:
116, pi. 7, figs. 20, 21, S, 9. Roewer, 1954,
Katalog der Araneae, 2: 530. Bonnet, 1955,
Bibliographia Araneorum 2: 349.
Diagnosis. Wulfila alba is closest to W.
sahaJninda but may be distinguished from
it by its lack of dark markings, the .spur-
like retrolateral tibial apophysis (Fig. 84)
and the shape of the spermathecae (Fig.
100).
Male (Orange Co., Florida). Coloration
as in Wulfila saltahunda except that dark
markings are entirely absent.
Total length 3.65 mm. Carapace 1.57
mm long, 1.21 mm wide, cephalic width
0.59 mm, clypeus height 0.07 mm. Eyes:
diameters (mm): AME 0.04, ALE 0.07,
PME 0.07, PLE 0.07; anterior eye row 0.36
mm long, straight; posterior eye row 0.49
mm long, procurved; MOQ length 0.22
mm, front width 0.12 mm, back width
246 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
0.26 mm; eye interdistances (mm): AME-
AME 0.04, AME-ALE 0.05, PME-PME
0.11, PiME-PLE 0.06, ALE-PLE 0.04.
Sternum 0.95 mm long, 0.70 mm wide.
Chelicerae 0.45 mm long with 6 promar-
ginal teeth and 7 retromarginal denticles.
Abdomen 2.12 mm long, 1.15 mm wide.
Epigastric furrow 0.67 mm from tracheal
spiracle, spiracle 0.79 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 3.13,
4; II 1.85, 7; III 1.12, 13; IV 2.03, 6. Ven-
tral spination: tibiae I 2-2-0, II 1-2-0, III
0-1-0, IV 1-1-0; metatarsi I, II, III 2-2-0,
IV 1-1-2.
Palpus as in Figures 83, 84.
Female (Indian River Co., Florida).
Coloration as in male.
Total length 4.00 mm. Carapace 1.62
mm long, 1.28 mm wide; cephalic width
0.58 mm, clypeus height 0.06 mm. Eyes:
diameters (mm): AME 0.04, ALE 0.07,
PME 0.08, PLE 0.08; anterior eye row 0.40
mm long, straight; posterior eye row 0.54
mm long, procurved; MOQ length 0.25 mm,
front width 0.14 mm, back width 0.29 mm;
eye interdistances (mm): AME-AME
0.05, AME-ALE 0.06, PME-PME 0.13,
PME-PLE 0.10, ALE-PLE 0.06.
Sternum 0.92 mm long, 0.74 mm wide.
Chelicerae 0.41 mm long with 6 promar-
ginal teeth and 9 retromarginal denticles.
Abdomen 2.66 mm long, 1.51 mm wide.
Epigastric furrow 0.68 mm from tracheal
spiracle, spiracle 0.88 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 3.13,
4; II 1.91, 8; III 1.05, 13; IV 2.07, 8. Ven-
tral spination as in male except tibiae II
2-2-0 and III 1-2-0 and metatarsi IV 2-
2-2.
Epigynum as in Figure 90, internal geni-
talia as in Figure 100.
Natural history. Mature males have been
taken from late March through early Au-
gust, mature females from early April
through late August. Specimens have been
taken by sweeping, on pines, and in Ma-
laise and pitfall traps.
Distribution. Maryland west to southern
Illinois, south to Florida and eastern Texas
(Map3).
Wulfila tantilla Chickering
Map 3; Figures 85, 86, 91, 101
Cragus palUdus O. P. -Cambridge, 1896, Biologia
Centrali Americana, Aran., 1: 215, pi. 26, fig.
10 { $). Male holotype from Santa Ana, Guate-
mala, in BMNH, examined. Preoccupied by
Wulfila pallidus O. P. -Cambridge, 1895. Roewer,
1954, Katalog der Araneae, 2: 535. Bonnet,
1956, Bibliographia Araneorum, 2: 1246.
Wulfila tantilla Chickering, 1940, Trans. Amer.
Microsc. Soc, 59: 119, figs. 64-66 {$). Male
holotype from El Valle, Panama, in MCZ,
examined. Roewer, 1954, Katalog der Araneae,
2: 555. NEW SYNONYMY.
Wulfila tenella Chickering, 1940, Trans. Amer.
Microsc. Soc, 59: 120, figs. 67, 68 ( $ ). Fe-
male holotype from El Valle, Panama, in MCZ,
examined. Roewer, 1954, Katalog der Araneae,
2: 555. NEW SYNONYMY.
Diagnosis. Wulfila tantilla is very closely
related to W. irnmaculella but may be dis-
tinguished by the greath' expanded tip of
the retrolateral tibial apophysis (Fig. 86)
and by the epigynal ducts terminating far
anterior of the epigynal openings (Fig.
91).
Male (Webb Co., Texas): Coloration as
in Wulfila alba, except that the posterior
declivity of the carapace is darkened.
Total length 3.02 mm. Carapace 1.62
mm long, 1.12 mm wide, cephalic width
0.63 mm, clypeus height 0.09 mm. Eyes:
Plate 8
Figures 92, 94, 107. Left palpi, ventral view. Figures 93, 95, 105. Left palpi, retrolateral view. Figures 96-98,
106. Epigyna, ventral view. Figures 99-104, 108. Internal genitalia, dorsal view. Figure 109. Body, dorsal view.
92, 93, 96, 102. Wulfila bryantae new species. 94, 95, 97, 104. Wulfila wunda new species. 98, 103. Wulfila
irnmaculella (Gertsch). 99. Wulfila saltabunda (Hentz). 100. Wulfila alba (Hentz). 101. Wulfila tantilla Chick-
ering. 105-109. Oxysoma cubana Banl<s. (Figs. 105, 106, 109 by Wilton Ivie, not to scale.)
Spider Family Anyphaenidae • Plalnick 247
248 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
diameters (mm): AME 0.06, ALE 0.07,
PME 0.09, PLE 0.09; anterior eye row
0.41 mm long, straight; posterior eye row
0.53 mm long, procurved; MOQ length
0.23 mm, front width 0.16 mm, back width
0.30 mm; eye interdistances (mm): AME-
AME 0.05, AME-ALE 0.04, PME-PME
0.13, PME-PLE 0.06, ALE-PLE 0.04.
Sternum 0.74 mm long, 0.63 mm wide.
Chelicerae 0.58 mm long with 5 promar-
ginal teeth on a carina and 8 retromarginal
denticles.
Abdomen 1.57 mm long, 0.97 mm wide.
Epigastric furrow 0.50 mm from tracheal
spiracle, spiracle 0.44 mm from base of
spinnerets.
Coxae III and IV with two small knobs.
Tibial lengths (mm) and indices: I 2.77, 3;
II 1.51, 9; III 0.86, 15; IV 1.51, 9. Ventral
spination: tibiae I, II 2-2-0, III, IV 1-2-0;
metatarsi I, II 2-2-0, III, IV 2-2-2.
Palpus as in Figures 85, 86.
Female (Hidalgo Co., Texas). Colora-
tion as in male of Wulfila alha.
Total length 2.92 mm. Carapace 1.34
mm long, 0.99 mm wide, cephalic width
0.67 mm, clypeus height 0.08 mm. Eyes:
diameters (mm): AME 0.05, ALE 0.06,
PME 0.06, PLE 0.06; anterior eye row 0.37
mm long, straight; posterior eye row 0.50
mm long, procurved; MOQ length 0.20
mm, front width 0.16 mm, back width 0.26
mm; eye interdistances ( mm ) : AME-
AME 0.05, AME-ALE 0.04, PME-PME
0.13, PME-PLE 0.06, ALE-PLE 0.05.
Sternum 0.89 mm long, 0.61 mm wide.
Chelicerae 0.50 mm long with 4 promar-
ginal teeth and 6 retromarginal denticles.
Abdomen 1.62 mm long, 1.15 mm wide.
Epigastric furrow 0.59 mm from tracheal
spiracle, spiracle 0.52 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 2.36, 5; II 1.21, 11; III 0.77,
18; IV 1.40, 11. Ventral spination as in
male save tibiae IV 1-1-0.
Epigynum as in Figure 91, internal geni-
talia as in Figure 101.
Natural history. Mature males have been
taken from mid-April through mid-Octo-
ber, mature females apparently year-round.
Nothing is known of the habits of this
species.
Distribution. Southern Texas south to
the Canal Zone (Map 3).
Wulfila immaculella (Gertsch),
new combination
Map 3; Figures 87, 88, 98, 103
Amjphac'uella immaculella Gertsch, 1933, Amer.
Mus. Novitates, No. 637: 9, fig. 14 ( ? ). Fe-
male holotype from Sabino Basin, Santa Cata-
lina Movmtains, Arizona, in AMNH, examined.
Roewer, 1954, Katalog der Araneae, 2: 530.
Bonnet, 1955, Bibliographia Araneorum, 2: 349.
Diagnosis. Wulfila immaculella is very
closely related to W. tantilla but may be
distinguished by the unexpanded tip of the
retrolateral tibial apophysis (Fig. 88) and
by the epigynal ducts terminating near the
epigynal openings (Fig. 98).
Male (Sonora, Mexico). Coloration as
in Wulfila alba.
Total length 3.60 mm. Carapace 1.64
mm long, 1.12 mm wide, cephalic width
0.67 mm, clypeus height 0.08 mm. Eyes:
diameters (mm): AME 0.05, ALE 0.08,
PME 0.08, PLE 0.08; anterior eye row 0.40
mm long, straight; posterior eye row 0.52
mm long, procurved; MOQ length 0.22
mm, front width 0.14 mm, back width
0.31 mm; eye interdistances (mm): AME-
AME 0.05, AME-ALE 0.05, PME-PME
0.14, PME-PLE 0.08, ALE-PLE 0.04.
Sternum 0.90 mm long, 0.68 mm wide.
Chelicerae 0.51 mm long with 4 promar-
ginal teeth and 5 retromarginal denticles.
Abdomen 2.07 mm long, 1.00 mm wide.
Epigastric furrow 0.63 mm from tracheal
spiracle, spiracle 0.74 mm from base of
spinnerets.
Coxae III with one, coxae IV with two
small knobs. Tibial lengths (mm) and in-
dices: I 3.42, 3; II 2.05, 5; III 1.30, 11; IV
2.11, 8. Ventral spination: tibiae I 2-2-0,
II 1-2-0, III 0-1-0, IV 1-1-0; metatarsi I,
II 2-2-0, III, IV 2-1-2.
Palpus as in Figures 87, 88.
Spider Family Anyphaenidae • Plat nick 249
Fenmle (Sonora, Mexico). Coloration as
in male of Wiilfila alba.
Total length 3.64 mm. Carapace 1.5S
mm long, 1.08 mm wide, cephalic width
0.61 mm, clypen.s height 0.10 mm. Eyes:
diameters (mm): AME 0.05, ALE 0.07,
PME 0.08, PLE 0.07; anterior eye row
0.40 mm long, straight; posterior eye row
0.53 mm long, procurved; MOQ length
0.22 mm, front width 0.15 mm, back width
0.30 mm; eye interdistances (mm): AME-
AiME 0.05, AME-ALE 0.04, PME-PME
0.12, PME-PLE 0.08, ALE-PLE 0.04.
Sternum 0.94 mm long, 0.71 mm wide.
Chelicerae 0.53 mm long with teeth as in
male.
Abdomen 2.05 mm long, 1.40 mm wide.
Epigastric furrow 0.65 mm from tracheal
spiracle, spiracle 0.74 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 2.81, 4; II 1.62, 9; III 0.92,
15; IV 1.75, 8. Ventral spination as in male
except tibiae II 2-2-0 and III 1-1-0.
Epigynum as in Figure 98, internal geni-
talia as in Figure 103.
Natural Jiistory. Mature males have been
taken in July, mature females in June and
July. One male was taken on Platanus sp.
Distrihuiion. Southern Arizona and So-
nora (Map 3).
Wulfila bryantae new species
Map 3; Figures 92, 93, 96, 102
Types. Male holotype, female paratype
from 5 miles east of Edinburg, Hidalgo
Co., Texas, 20 April 1937 (S. Mulaik), de-
posited in AMNH. Male and female para-
types from Jim Wells and Cameron Coun-
ties, Texas, deposited in MCZ. The
specific name is a patronym in honor of
Miss Elizabeth Bryant, in recognition of her
pioneering work on North American any-
phaenids.
Diagnosis. Wulfila bryantae is a distinc-
tive species easily recognized by its stubby
median apophysis (Fig. 92) and the medial
ridge on the epigynum (Fig. 96).
Male (Hidalgo Co., Texas). Coloration
as in Wulfila alba.
Total length 3.35 mm. (Carapace 1.44
mm long, 1.08 mm wide, cephalic width
0.81 mm, clypeus height 0.10 mm. Eyes:
diameters (mm): AME 0.06, ALE 0.06,
PME 0.07, PLE 0.07; ant(>rior eye row 0.49
mm long, slightly rcx-urved; posterior eye
row 0.62 mm long, procurved; MOQ length
0.26 mm, front width 0.20 mm, back width
0.30 mm; eye interdistances (mm): AME-
AME 0.07,^ AME-ALE 0.07, PME-PME
0.15, PME-PLE 0.13, ALE-PME 0.06.
Sternum 0.97 mm long, 0.55 mm wide.
Chelicerae 0.73 mm long with 3 promar-
ginal teeth on a carina and 7 retromarginal
denticles.
Abdomen 1.80 mm long, 1.12 mm wide.
Epigastric fvnrow 0.56 mm from tracheal
spiracle, spiracle 0.68 mm from base of
spinnerets.
Coxae I with a small knob, coxae II with
two spurs. Tibial lengths (mm) and in-
dices: I 2.76, 4; II 1.85, 7; III 0.92, 15; IV
1.89, 7. \'entral spination: tibiae I, II 2-
2-0, III 1-2-0, IV 1-1-0; metatarsi I, II 2-
2-0, III, IV 2-1-2.
Palpus as in Figures 92, 93.
Female (Hidalgo Co., Texas). Colora-
tion as in male of Wulfila alba.
Total length 3.78 mm. Carapace 1.44
mm long, 0.99 mm wide, cephalic width
0.74 mm, clypeus height 0.07 mm. Eves:
diameters (mm): AME 0.06, ALE 6.07,
PME 0.06, PLE 0.07; anterior eye row
0.42 mm long, slightly recurved; posterior
eye row 0.59 mm long, procurved; MOQ
length 0.23 mm, front width 0.17 mm, back
width 0.27 mm; eye interdistances (mm):
AME-AME 0.06, AME-ALE 0.05, PME-
PME 0.14, PME-PLE 0.13, ALE-PLE 0.06.
Sternum 0.74 mm long, 0.64 mm wide.
Chelicerae 0.62 mm long with 5 promar-
ginal teeth and 5 retromarginal denticles.
Abdomen 2.59 mm long, 2.16 nun wide.
Epigastric furrow 0.90 mm from tracheal
spiracle, spiracle 0.88 mm from base of
spinnerets.
250 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
Legs unmodified. Tibial lengths (mm)
and indices: I 2.24, 5; II 1.37, 9; III 0.72,
19; IV 1.35, 10. Ventral spination as in
male except tibiae III, IV 2-2-0 and meta-
tarsi III, IV 2-0-2.
Epigynum as in Figure 96, internal
genitalia as in Figure 102.
Natural history. Mature males have been
taken from late April through early June,
mature females from early April through
early December. Nothing is known of the
habits of this species.
Distribution. Southern Texas and Ta-
maulipas (Map 3).
Wulfila wunda new species
Map 3; Figures 94, 95, 97, 104
Wulfila immaculata, Bryant (not Banks), 1936,
Psyche, 43: 98, fig. 1, $. Male allotype from
Brichell Hammock, Florida Keys, in MCZ,
examined. Not Wulfila immaculata Banks, 1914,
Bull. Amer. Mus. Natur. Hist., 33: 640, pi. 43,
fig. 7, 9 . Female holotype from Vinales, Pinar
del Rio, Cuba, in AMNH, examined.
Types. Male holotype, female paratype
from Tavernier, Monroe Co., Florida, 16
February 1951 (A. M. Nadler), deposited
in AMNH. Male and female paratypes
from Dade Co., Florida, deposited in MCZ.
The specific name is an arbitrary combina-
tion of letters.
Diagnosis. Wulfila wunda is a distinc-
tive species the genitalia of which are
quite different from those of the other
Wulfila in America north of Mexico: the
retrolateral tibial apophysis is very short
( Fig. 95 ) and the epigynum has an atrium
(Fig. 97).
Male (Dade Co., Florida). Coloration
as in Wulfila alba.
Total length 3.42 mm. Carapace 1.55
mm long, 1.08 mm wide, cephalic width
0.68 mm, clypeus height 0.06 mm. Eyes:
diameters (mm): AME 0.05, ALE 0.07,
PME 0.08, PLE 0.08; anterior eye row 0.48
mm long, straight; posterior eye row 0.59
mm long, procurved; MOQ length 0.20
mm, front width 0.14 mm, back width 0.29
mm; eye interdistances (mm): AME-
AME 0.05, AME-ALE 0.09, PME-PME
0.14, PME-PLE 0.12, ALE-PLE 0.04.
Sternum 1.06 mm long, 0.70 mm wide.
Chelicerae 0.85 mm long with 4 promar-
ginal teeth and 6 retromarginal denticles.
Abdomen 1.91 mm long, 1.01 mm wide.
Epigastric furrow 0.70 mm from ti'acheal
spiracle, spiracle 0.76 mm from base of
spinnerets.
Coxae II with a small knob. Tibial
lengths (mm) and indices: I 4.10, 3; II
1.87, 8; III 1.01, 15; IV 2.05, 7. Ventral
spination: tibiae I, II 2-2-0, III 0-1-0, IV
0-2-0; metatarsi I, II 2-2-0, III 0-2-0, IV
2-1-2.
Palpus as in Figures 94, 95.
Female (Dade Co., Florida). Colora-
tion as in male of Wulfila alba.
Total length 3.74 mm. Carapace 1.55
mm long, 1.15 mm wide, cephalic width
0.72 mm, clypeus height 0.07 mm. Eyes:
diameters (mm): AME 0.04, ALE 0.06,
PME 0.07, PLE 0.07; anterior eye row 0.49
mm long, straight; posterior eye row 0.59
mm long, procurved; MOQ length 0.20
mm, front width 0.15 mm, back width
0.30 mm; eye interdistances ( mm ) : AME-
AME 0.06, AME-ALE 0.10, PME-PME
0.16, PME-PLE 0.12, ALE-PME 0.04.
Sternum 0.90 mm long, 0.67 mm wide.
Chelicerae 0.65 mm long with 5 promar-
ginal teeth and 9 retromarginal denticles.
Abdomen 2.16 mm long, 1.15 mm wide.
Epigastric furrow 0.74 mm from tracheal
spiracle, spiracle 0.83 mm from base of
spinnerets.
Legs unmodified. Tibial lengths (mm)
and indices: I 3.13, 4; II 1.44, 10; III 0.76,
20; IV 1.58, 9. Ventral spination as in male
except tibiae III 1-2-0 and IV 0-1-0 and
metatarsi III 1-2-0 and IV 1-2-2.
Epigynum as in Figure 97, internal geni-
talia as in Figure 104.
Natural history. Mature males have been
taken from mid-February through mid-
May, mature females apparently year-
round. Nothing is known of the habits of
this .species.
Spider Family Anyphaenidae • Platnick 251
Distrihuiion. Southern Florida, Culxi,
and Mona Island ( Map 3).
Aysha Keyserling
Aysha Keyserling, 1891, Spinn. Ainer. ( Brasil.
Spiiin.), 3: 83, 129. Type species Aysha pros-
pera Keyserling, 1891, designated by Simon,
1897, Hist. Natm-. Araig., 2: 104.
Diagnosis. Aysha is easily recognized by
the greatly adxanced placement of the
tracheal spiracle, located just behind the
epigastric furrow. The genitalic structure
is quite different from that of Amjphacna
and Wulfila and the genus undoubtedly
represents a different evolutionary line.
There are probably more than thirty spe-
cies in this genus; they occur commonly
in both North and South America.
Description. Total length 4-9 mm. Cara-
pace longer than wide, narrowed in front
to more than half its maximum width.
Clypeus height roughly equal to anterior
median eye diameter. All eyes subequal in
size. Procurved posterior eye row longer
than recurved anterior row. Median ocular
quadrangle longer than wide in front,
wider in back than long. Anterior median
eyes separated by slightly less than their
diameter, slightly closer to anterior laterals.
Posterior medians separated by up to twice
their diameter, closer to posterior laterals.
Anterior laterals separated by their radius
from posterior laterals. Sternum longer
than wide, unmodified. Chelicerae with 3-
4 promarginal teeth and 7-9 retromarginal
denticles. Abdomen longer than wide,
tracheal spiracle much closer to epigastric
furrow than to base of spinnerets. Leg
formula 1423, legs unmodified. Metatarsi I
and II with one pair of \'entral spines. Pal-
pus with greatly enlarged base of embolus,
long curving embolus and short conductor.
Ventral tibial apophysis sometimes present
in addition to retrolateral tibial apophysis.
Epigynum with anterior median opening
and two sidepieces. Internal genitalia with
long, sometimes coiling, ducts.
Variation. Only Aysha gracilis shows
significant variation, and that is in size and
not strnetine or coloration. The size of both
the whole animal and ol the genitalia vary
geographically. The largest .specimens
( males with cymbium length averaging 1.3
mm) occur in Virginia and surrounding
states, with smaller individuals occurring
in the north (New England and Michigan
males with cymbium length averaging 1.1
mm) and in the south (Texas males with
cymbium length averaging 0.9 mm).
Key to Species
la. Males 2
lb. Females -..- 7
2a. Palpus without a ventral tibial apophysis
(VTA) (Figs. Ill, 119) -.._ 3
2b. Palpus with a ventral tibial apophysis
(VTA), sometimes small, transparent, eas-
ily overlooked (Figs. 113, 115, 117, 121) 4
3a. Embolus restricted to distal half of palpal
bulb (Fig. 118) arunda
3b. Embolus not restricted to distal half of pal-
pal bulb (Fig. 110) velox
4a. VTA erect, sclerotized, relatively large
(Figs. 113, 115) 5
4b. VTA recumbent, transparent, relativelv
small (Figs. 117, 121) '. 6
5a. Distal retrolateral tip of tegulum with a
flap covering embolus (Fig. 112) decepta
5b. Distal retrolateral tip of tegulum with a
sharp point underlying embolus (Fig. 114)
— incursa
6a. Base of embolus recurved, with a sharp
spike (Fig. 120) camhridgei
6b. Base of embolus not recurved, forming a
smooth arc (Fig. 116) fitacilis
7a. Internal genitalia with simple uncoiled
ducts (Figs. 124, 127, 141, 143) 9
7b. Internal genitalia coiled or with accessory
ducts (Figs. 125, 142) 8
8a. Internal genitalia highly coiled ( Fig.
125 ) '...... vclox
8b. Internal genitalia not coiled but with loop-
ing accessory ducts (Fig. 142) arunda
9a. Median epig\'nal opening near anterior
rim (Figs. 123, 126, 138) 10
91). Median epigynal opening near middle of
epigynum ( Fig. 140 ) -.. gracilis
10a. Median epig\nal opening much wider than
epigynal sidepieces (Fig. 138) cand)ridgc'i
lOb. Median epigynal opening not wider than
epigynal sidepieces (Figs. 123, 126) 11
11a. Base of epig>nal sidepieces near epigastric
furrow (Fig. 126); internal genitalia with
angular ducts (Fig. 127) incursa
252 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
. ^ Aysha gracilis
I /
; 1 / HM
~T— 7
I
Ayshc velox
r?
i?
♦^^'^r
cQ^*
i
i
" 1 \
r'
)
(
-A
Aysha arunda
.„ V
Map 4. Distributions of Aysha arunda. A. cambridgei, A. decepta, A. gracilis, A. incursa and A. velox.
111). Base of epigynal sidepieces far from
epigastric furrow (Fig. 123); internal
genitalia with rounded ducts (Fig. 124)
decepta
Aysha prospers Keyserling
Figure 145
Ay.sha prospera Keyserling, 1891, Spinnen Amer-
ikas (Brasil. Spinn.), 3: 129, pi. 4, fig. 88
( 9 ). Female holotype from Rio Grande, Brasil,
in BMNH, examined. Roewer, 1954, Katalog
der Araneae, 2: 533. Bonnet, 1955, Biblio-
graphia Araneorum, 2: 838.
This South American species, type spe-
cies of Aysha, is a member of a large, dis-
tinct species group. Somatic characters
clearly ally it with the North American
forms included in the genus.
Aysha gracilis (Hentz)
Map 4; Figures 116, 117, 140, 143
Chihiona gracilis Hentz, 1847, J. Boston Soc.
Natur. Hist., 5: 452, pi. 23, fig. 9(5). Type
specimens from North Carolina and Alabama in
Boston Soc. Natur. Hist. (Boston Museum of
Science), destroyed by beetles.
Anijphaena gracilis, L. Koch, 1836, Arach. Fam.
brassidae, p. 195, pi. 8, fig. 130, 9 . Comstock,
1912, Spider Book, p. 561, fig. 633, $ (not
fig. 632).
Anijphaena rubra Emerton, 1890, Trans. Connecti-
cut Acad. Sci., 8: 186, pi. 6, fig. 1(9). Male
allotype (?) from Franklin Park, Boston, Mas-
sachusetts, in MCZ, examined. Emerton, 1909,
Trans. Connecticut Acad. Sci., 14: 220, pi. 9,
fig. 8-8c, $ .
Aysha gracilis, Bryant, 1931, Psyche, 38: 119, pi.
7, fig. 13, pi. 8, fig. 26, $, 9. Chickering,
1939, Pap. Michigan Acad. Sci., 24: 53, figs.
9-11, $, 9. Comstock, 1940, Spider Book,
rev. ed., p. 575, fig. 633, $ (not fig. 632).
Kaston, 1948, Bull. Connecticut Geol. Natur.
Hist. Surv., 70: 405, figs. 1452, 1459-1464, $,
9 . Roewer, 1954, Katalog der Araneae, 2: 534.
Bonnet, 1955, Bibliographia Araneorum, 2: 837.
Diagnosis. Aysha gracilis is closest to
A. cambridgei but lacks the sharp spike on
the proximal edge of the base of embolus
(Fig. 116) of that .species. Females have
Spider Family Anyphaenidae • Platnick 253
Plate 9
Figures 110, 112, 114, 116. Left palpi, ventral view. Figures 111, 113, 115, 117
110, 111. Aysha velox (Becker). 112, 113. Aysha decepta (Banks)
116, 117. Aysha gracilis (Hentz).
Left palpi, retrolateral view/.
114, 115. Aysha incursa (Chamberlin).
the median epigynal opening near the mid- long, 2.02 mm wide, cephalic width 1.17
die of the epigynum (Fig. 140). Variation mm, clypeus heigiit 0.09 mm, light orange-
in thi.s species is discussed above. brown, darkest anteriorly, with thin dark
Male (Middlesex Co., Massachusetts), border and two dark paramedian longitudi-
Total length 5.73 mm. Carapace 2.56 mm nal bands. Eyes: diameters (mm): AME
254
Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
0.09, ALE 0.11, PME 0.09, PLE 0.11; an-
terior eye row 0.60 mm long, slightly re-
cnrved; posterior eye row 0.80 mm long,
procurved; MOQ length 0.32 mm, front
width 0.26 mm, back width 0.38 mm; eye
interdistances (mm): AME-AME 0.09,
AME-ALE 0.07, PME-PME 0.19, PME-
PLE 0.15, ALE-PLE 0.06.
Sternum 1.44 mm long, 1.08 mm wide,
light orange-brown with translucent border
and darkened extensions to coxae. Chelic-
erae 1.12 mm long with 4 promarginal
teeth and 8 retromarginal denticles, dark
orange-brown proximally, dark brown dis-
tally. Labium and endites light orange-
brown, darkest proximally. Endites sharply
invaginated at middle.
Abdomen 3.20 mm long, 1.73 mm wide,
pale grayish-brown with transverse rows
of dark markings, venter pale. Epigastric
furrow 0.40 mm from tracheal spiracle,
spiracle 1.73 mm from base of spinnerets.
Legs light orange-brown with distal seg-
ments darkest. Tibial lengths (mm) and
indices: I 2.64, 10; II 1.87, 15; III 1.19, 26;
IV 2.09, 15. Vential spination: tibiae I, II
2-2-2, III 1-2-2; IV 2-2-2; metatarsi I, II
2-0-0, III 2-1-2, IV 2-2-2.
Palpus as in Figure 116, 117.
Female (Washington Co., Arkansas).
Coloration as in male.
Total length 8.42 mm. Carapace 2.75
mm long, 2.11 mm wide, cephalic width
1.47 mm, clypeus height 0.10 mm. Eyes:
diameters (mm): AME 0.14, ALE 0.14,
PME 0.13, PLE 0.14; anterior eye row 0.43
mm long, recurved; posterior eye row 1.04
mm long, procurved; MOQ length 0.43
mm, front width 0.36 mm, back width 0.49
mm; eye interdistances (mm): AME-
AME 0.09, AME-ALE 0.08, PME-PME
0.22, PME-PLE 0.18, ALE-PLE 0.06.
Sternum 1.84 mm long, 1.31 mm wide.
Chelicerae 1.57 mm long with teeth as in
male.
Abdomen 5.76 mm long, 3.53 mm wide.
Epigastric furrow 0.68 mm from tracheal
spiracle, spiracle 3.24 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 2.56,
16; II 1.94, 20; III 1.26, 30; IV 2.30, 17.
Ventral spination: tibiae I 2-2-0, II 1-2-1,
III 1-1-2, IV 1-2-2; metatarsi I, II 2-0-0,
III 2-0-2, IV 2-2-2.
Epigynum as in Figure 140, internal
genitalia as in Figure 143.
Natural history. Mature males and fe-
males have been taken year-round. Speci-
mens have been taken by sweeping, in
pitcher plants, on loblolly pine, in fall web-
worm nests and frequently in houses.
Distribution. New England west to Wis-
consin and Iowa, south to Florida and east-
ern Texas ( Map 4 ) .
Aysha cam bridge! Bryant
Map 4; Figures 120, 121, 138, 141
Aysha cambiidgei Bryant, 1931, Psyche, 38: 119,
pi. 7, fig. 15 { $ ). Male holotype from
Guanajuato, Mexico, in MCZ, examined. Roe-
wer, 1954, Katalog der Araneae, 2: 532. Bon-
net, 1955, Bibliographia Araneorum, 2: 836.
Diagnosis. Aysha cambridgei is closely
related to A. gracilis but has a distinctive
spike on the proximal edge of the base of
the embolus (Fig. 120) and the median
epigynal opening near the anterior rim of
the epigynum ( Fig. 138 ) .
Male (Jeff Davis Co., Texas). Colora-
tion as in Aysha gracilis except that the ab-
domen is pale white with two dark para-
median longitudinal bands.
Total length 5.87 mm. Carapace 2.41
mm long, 1.91 mm wide, cephalic width
0.97 mm, clypeus height 0.11 mm. Eyes:
diameters (mm): AME 0.11, ALE 0.12,
PME 0.11, PLE 0.11; anterior eye row 0.57
mm long, recurved; posterior eye row 0.75
mm long, procurved; MOQ length 0.33
mm, front width 0.28 mm, back width 0.38
mm; eye interdistances (mm): AME-
AME 0.06, AME-ALE 0.05, PME-PME
0.16, PME-PLE 0.11, ALE-PLE 0.05.
Sternum 1.42 mm long, 1.01 mm wide.
Chelicerae 0.98 mm long with 4 promar-
ginal teeth and 7 retromarginal denticles.
Abdomen 3.49 mm long, 1.58 mm wide.
Epigastric furrow 0.68 mm from tracheal
Spideu Family Anyphaenidae • Platnick 255
122
126
127
125
Plate 10
Figures 118, 120. Left palpi, ventral view. Figures 119, 121. Left palpi, retrolateral view. Figures 122, 123,
126. Epigyna, ventral view. Figures 124, 125, 127. Internal genitalia, dorsal view. 118, 119. Aysha arunda new
species. 120, 121. Aysha cambridgei Bryant. 122, 125. Aysha velox (Becker). 123, 124. Aysha decepta
(Banks). 126, 127. Aysha incursa (Chamberlin).
.spiracle, spiracle 1.55 mm from base ot Ventral spination: tibiae I, II 2-2-2, III
spinnerets. 1-2-2, IV 2-2-2; metatarsi I, II 2-0-0, III,
Tibial lengths (mm) and indices: I 3.06, IV 2-2-2.
8; II 1.87, 13; III 1.28, 21; IV 2.16, 12. Palpus as in Figures 120, 121.
256 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4
Female (Henderson Co., Texas). Color-
ation as in male.
Total length 8.50 mm. Carapace 3.35
mm long, 2.52 mm wide, cephalic width
1.69 mm, clypeus height 0.12 mm. Eyes:
diameters (mm): AME 0.14, ALE 0.16,
PME 0.14, PLE 0.14; anterior eye row 0.84
mm long, recm'ved; posterior eye row 1.11
mm long, procurved; MOQ length 0.42
mm, front width 0.37 mm, back width 0.50
mm; eye interdistances (mm): AME-
AME 0.10, AME-ALE 0.07, PME-PME
0.22, PME-PLE 0.20, ALE-PLE 0.05.
Sternum 1.91 mm long, 1.22 mm wide.
'&'
Chelicerae 1.69 mm long with teeth as in
male.
Abdomen 5.04 mm long, 2.88 mm wide.
Epigastric furrow 0.61 mm from tracheal
spiracle, spiracle 3.17 mm from base of
spinnerets.
Tibial lengths (mm) and indices: 12.88,
12; II 2.07, 17; III 1.40, 26; IV 2.57, 15.
Ventral spination as in male except tibiae
I, II 2-2-0 and III 2-2-2.
Epigynum as in Figure 138, internal
genitalia as in Figure 141.
Natural history. Mature males have been
taken from mid-June through early August,
mature females from late May through
early August. Specimens have been taken
on trees and shrubs.
Distribution. South central states from
Alabama to western Texas, south to central
Mexico (Map 4).
Aysha decepta (Banks)
Map 4; Figures 112, 113, 123, 124
Amjphaena decepta Banks, 1899, Proc. Ent. Soc.
Washington, 4: 190. Female holotype from
Brazos Co., Texas, in MCZ, examined.
Aysha mimita F. O. P.-Canibridge, 1900, Biologia
Centrali Americana, Aran., 2: 99, pi. 7, figs.
18-19 { $, ? ). Male holotype, female allotype
from Guatemala, in BMNH, examined. Bryant,
1931, Psyche, 38: 120, pi. 7, fig. 17, $. Roe-
wer, 1954, Katalog der Araneae, 2: 533. Bon-
net, 1955, Bibliographia Araneorum, 2: 838.
NEW SYNONYMY.
Aijsha decepta, Bryant, 1931, Psyche, 38: 120, pi.
7, fig. 16, pi. 8, fig. 27, $, 9. Roewer, 1954,
Katalog der Araneae, 2: 534. Bonnet, 1955,
Bibliographia Araneorum, 2: 836.
Diagnosis. Aysha decepta is very closely
related to A. incursa but has a characteris-
tic flap (on the retrolateral tip of the tegu-
lum) that covers the embolus (Fig. 112),
while the base of the epigynal sidepieces
is a considerable distance from the epigas-
tric furrow ( Fig. 123 ) . Both morphological
and zoogeographical data (Map 4) indi-
cate that these two species are each other's
nearest relatives.
Male (Hidalgo Co., Texas). Coloration
as in Aysha camhridgei.
Total length 4.82 mm. Carapace 2.25
mm long, 1.76 mm wide, cephalic width
1.06 mm, clypeus height 0.10 mm. Eyes:
diameters (mm): AME 0.08, ALE 0.10,
PME 0.11, PLE 0.11; anterior eye row 0.58
mm long, straight; posterior eye row 0.75
mm long, procurved; MOQ length 0.23
mm, front width 0.24 mm, back width 0.39
mm; eye interdistances ( mm ) : AME-
AME 0.08, AME-ALE 0.06, PME-PME
0.18, PME-PLE 0.12, ALE-PLE 0.05.
Sternum 1.37 mm long, 0.85 mm wide.
Chelicerae 0.97 mm long with 4 promar-
ginal teeth and 7 retromarginal denticles.
Abdomen 2.74 mm long, 1.39 mm wide.
Epigastric furrow 0.38 mm from tracheal
spiracle, spiracle 1.28 mm from base of
spinnerets.
Tibial lengths (mm) and indices: I 2.54,
9; II 1.67, 14; III 1.01, 25; IV 1.89, 16.
Venti-al spination: tibiae I 2-2-0, II 1-2-0,
III 1-2-2, IV 2-2-2; m